/^BERKELEY* 

LIBRARY 

I     UNIVERSITY  OF 
\CAUFORNIA 


EARTH 

SCIENCES 

LIBRARY 


Cambridge  (geographical 

GENERAL  EDITOR  :   F.  H.  H.  GUILLEMARD,  M.D., 

FORMERLY   LECTURER   IN    GEOGRAPHY   IN  THE  UNIVERSITY  OF  CAMBRIDGE. 


A    GEOGRAPHICAL 
HISTORY    OF    MAMMALS. 


ilontion:    C.    J.    CLAY   AND   SONS, 
CAMBRIDGE   UNIVERSITY    PRESS   WAREHOUSE, 

AVE  MARIA   LANE. 
ffilaggofo:    263,  ARGYLE   STREET. 


:   F.  A.  BROCKHAUS. 
lorfe:   MACMILLAN  AND  CO. 


THE  WORLD  ON  MERCATOR'S  PROJECTION  SHEWING  THE 


Cambridge  TJniversi 


UVIMALIAN  GEOGRAPHICAL  REALMS  &  REGIONS. 


90-  ins-  i»o' 


London^:  Stamfords  Geog*'  Estab*. 


A    GEOGRAPHICAL 


HISTORY  OF  MAMMALS, 


BY 


R.    LYDEKKER,   B.A.,    F.R.S.,   V.P.G.S.  ETC. 


CAMBRIDGE: 

AT   THE   UNIVERSITY   PRESS. 
1896 

[All  Rig/its  reserved.} 


PALEONTOLOGY  LIBRARY 
Gift  of  C.  A.  Kofoid 


(JEamfartoge : 

PRINTED   BY  J.    AND   C.    F.    CLAY, 
AT   THE   UNIVERSITY  PRESS. 


L3 


PREFACE. 

SCIENCES 
LIBRARY 


SINCE  the  publication  of  Dr  Wallace's  book  on  the  geo- 
graphical distribution  of  animals  in  general,  the  only  works  which 
have  appeared  relating  to  Mammals  from  the  same  point  of  view 
are  the  small  volume  by  Mr  F.  E.  Beddard  and  the  series  of 
papers  by  Mr  W.  L.  Sclater,  referred  to  in  the  Appendix. 

Both  the  latter  admittedly  take  but  little  account  of  fossil 
forms ;  and  there  is  accordingly  ample  room  for  a  work  which 
should  collect  and  arrange  the  information  on  this  subject,  and 
indicate  the  deductions  which  may  be  drawn  therefrom.  This 
task  has  been  attempted  in  the  present  volume.  The  subject 
is,  however,  so  vast,  and  the  information  relating  to  it  scattered 
through  so  many  publications,  that  it  is  probable  many  points  of 
interest  or  importance  have  not  been  noticed.  From  the  mode 
of  arrangement  of  the  work,  a  considerable  amount  of  repetition 
was  inevitable. 

The  long  time  that  the  volume  was  in  the  press  will  account 
for  its  containing  no  allusion  to  certain  papers  which  have  ap- 
peared recently.  It  may  therefore  be  mentioned  here  that 
recent  discoveries  have  shown  a  slight  intermixture  of  northern 
types  in  the  Palaeozoic  flora  of  Argentina,  so  that  the  isolation 
of  the  northern  and  southern  floras  is  .not  so  nearly  complete 
as  was  supposed.  A  paper  just  published  by  M.  Boule1  indi- 

1  Comptes  RenduS)  vol.  cxxii  (1896). 


VI  PREFACE. 

cates  that  the  relationship  between  Cadurcotherium  and  the 
Astrapotheria  is  closer  than  suggested  on  page  82.  Hence  there 
is  further  evidence  that  South  America  received  its  Tertiary 
ungulates  by  way  of  Africa.  The  extinct  Patagonian  birds 
Phororhachis  and  Brontornis  appear,  according  to  Mr  C.  W. 
Andrews,  to  be  nearly  allied  to  the  existing  South  American 
seriema  (Cariama),  and  since  Filholornis,  of  the  French  Phos- 
phorites, has  been  shown  by  Prof.  A.  Milne-Edwards  to  be 
related  to  the  hoatzin  (Opisthocomus),  it  would  seem  that  all 
these  peculiar  South  American  types  of  birds  have  a  history 
somewhat  similar  to  that  of  the  extinct  ungulates  of  the  same 
region. 

R.    LYDEKKER. 

HARPENDEN, 

June  ist,  1896. 


CONTENTS. 

CHAPTER   I. 

INTRODUCTORY. 

Distributional  Area  and  Station — Influence  of  Temperature — Humidity — 
Other  Factors  in  Distribution — Importance  of  Palaeontology — Inequality 
in  the  Ages  of  different  Groups  of  Animals — Different  Groups  have  different 
Geographical  Distribution — Importance  of  Mammals  in  Geographical 
Distribution — Classification  of  Mammals — Barriers  to  Dispersal  of  Mam- 
mals— Influence  of  Man  on  Distribution — Extinction  of  the  larger 
Plistocene  Mammals — Distributional  Areas  of  Genera  and  Species — 
Centres  of  Evolution — Permanency  of  Continents  and  Ocean-Basins — 
Zoological  Realms  and  Regions  .......  i 

CHAPTER    II. 
THE  NOTOG^EIC   REALM. 

Definition  and  Characters  of  the  Realm — Australian  Region — Monotremes — 
Marsupials — Rodents — Carnivores — Ungulates — Bats — List  of  Australian 
and  Papuan  Genera — Polynesian  Region — Hawaiian  Region — Austro- 
Malayan  Region — Palseontological  History  of  Marsupials — How  Australia 
received  its  Fauna  .  .  .  .  .  .  .  .  .28 


CHAPTER   III. 

THE  NEOG^EIC  REALM. 

Extent  and  Characters — Mammaliferous  Deposits — Monkeys — Bats — Insecti- 
vora  —  Carnivores  —  Ungulates  —  Horses  — Litopterna — Astrapotheria — 
Toxodonts —  Pyrotheria — Proboscideans — Rodents  —  Edentates  —  Arma- 
dillos and  Glyptodonts — Sloths — Anteaters — Ground-sloths — Marsupials — 
Cetaceans — Early  Distinction  of  the  Neogseic  Fauna — Early  Separation 
of  N.  and  S.  America — Incursion  of  Northern  Mammals — Distinctness  of 
the  existing  Fauna — Origin  of  the  Santa  Cruz  Fauna — Antarctica  and  the 
South  American  element  in  the  Ethiopian  Fauna — Conclusion — Sub- 
regions 64 


Vlll  CONTENTS. 


CHAPTER    IV. 
THE  ARCTOG^EIC  REALM. 

Features  of  the  Arctogaeic  Fauna — Community  of  earliest  Fauna — Evidence  of 
Secondary  Reptiles — Puerco  Fauna — Lemuroids — Insectivora  —Carnivores 
— Rodents — Ungulates — Summary  of  the  Characteristics  of  the  Mamma- 
lian Fauna  of  Arctogaea  .  .  .  .  .  .  .  .  1 44 


CHAPTER   V. 

EASTERN  ARCTOG^EA. 

Mammalian  Groups  peculiar  to  Eastern  Arctogsea — Tertiary  Mammalian 
Fauna  of  Eastern  Arctogsea — Oligocene  Fauna — Miocene  Fauna — Older 
Pliocene  Fauna — Pikermi  and  allied  Faunas — Sivvalik  Fauna — Higher 
Pliocene  Faunas  .  .  .  .  .  .  .  .  .  •  1/9 


CHAPTER   VI. 

THE  MALAGASY   REGION. 

Mammalian  Fauna — Relations  of  Madagascar  to  the  Mainland         .         .     213 

CHAPTER   VII. 

THE   ETHIOPIAN   REGION. 

Characteristics  of  the  Mammalian  Fauna — Birds — Past  History  of  Ethiopia — 
Sub-regions  .  .  .  .  .  .  .  .  .  .  .227 

CHAPTER   VIII. 

THE   ORIENTAL   REGION. 

Sub-regions — Characteristics  of  the  Mammalian  Fauna — Past  History  of  the 
Region — Malayan  Sub-region — Nicobars,  Mentawi,  and  Christmas  Islands 
— Philippine  Sub-region  .........  264 


CONTENTS.  IX 


CHAPTER    IX. 

THE  HOLARCTIC   REGION. 

Characteristics  of  the  Mammalian  Fauna — Mammals  of  the  Eastern  Holarctic 
Region — Plistocene  Fauna  of  the  Holarctic  Region — Geographical  Changes 
since  the  Plistocene — Western  Division  of  the  Region — Arctic  Sub-region 
— European  Sub-region — Central  Asian  Sub-region — Tibetan  Sub-region 
— Mantchurian  Sub-region — Mediterranean  Sub-region — Kashmir — Ca- 
nadian Sub-region — Transition  Zone  .  .  .  .  .  .  308 


CHAPTER  X. 

THE  SONORAN   REGION. 

Limits — Characteristics  of  Mammalian  Fauna — Extinct  Groups  of  Mammals 
characteristic  of  Western  Arctogrea — Distinctness  of  the  Region — Dual 
origin  of  Groups  .  .  ..  .  .  .-..'.''  V  '•'  '.  363 


APPENDIX. 

LIST  OF  WORKS  AND  PAPERS      .-   &*&,      .        .        ..:..-,  fj       .    383 
INDEX    .        .        .        . V       .     386 


MAP   SHEWING   DISTRIBUTION   OF  ANIMALS  .  .      .    ,  to  face  Title 


LIST    OF    ILLUSTRATIONS. 

PAGE 

Fig.  i.  The  Duckbill  {Ornithorhyncus  anatinus]          .         .         .         .32 

„  2.     Skull  of  Rat-Kangaroo 35 

,,  3.  Skull  of  Extinct  Phalanger  ( Thylacoleo  carnifex)                               37 

„  4.  Banded  Anteater  (Myrmecobius  fasciatus]          ....       39 

,,  5.  Fore  Part  of  Skull  of  Babirusa  (Babirusa  alfums}    .         .         -47 

,,  6.  Lower  Jaw  of  Triconodon,  or  Allied  Form        .         .         .         .52 

,,  7.  Lower  Jaw  of  Phascolotheriiim           .         .         .         .         .         -53 

,,  8.  Lower  Jaw  of  Amphilestes        .         .         .         .         .         .         -53 

,,  9.  Lower  Jaw  of  Homunculus      .......       69 

,,  10.  Skeleton  of  Macrauchenia  patachonica     .....       76 

,,  n.     Skeleton  of  Hind  Foot  of  Rhinoceros 78 

,,  12.  Carpus  and  Metacarpus  of  Hyrax  and  Elephant        .         .         -78 

,,  13.     Upper  Molar  of  Macraiichenia .80 

,,  14.  Upper  Molar  of  Oxyodontotherium  ......       80 

,,  15.  Upper  Molar  of  Astrapotherium      .         .         .         .         .         .       81 

„  16.     Skull  of  Nesodon 83 

,,  17.  Palate  of  Typotherium     ........       84 

„  1 8.  External  Skeleton  of  Armadillo  ( Tatusia  giganted]  .         .         .       93 

„  19.  Internal  Skeleton  of  a  Glyptodont  ......       95 

,,  20.  External  Skeleton  of  Glyptodon  clavipes   .         .         .         .         -97 

,,  21.  External  Skeleton  of  Panochthus  tuberculatus .         .         .         -99 

,,  22.     Tamandua  Anteater 102 

,,  23.  Skull  of  Mylodon     .........      104 

,,  24.  Skeleton  of  Scelidothcrium  leptocephalum          .         .         .         .106 

„  25.  Lower  Jaw  of  Prothylacinus    .......     108 

,,  26.  Lower  Jaw  of  Acdestis  oweni    .         .         .         .         .         .         .no 

,,  27.  Lower  Jaw  of  Abderites  .         .         .         .         .         •         •         .     rii 

,,  28.     Lower  Jaw  of  Plagiaulax 149 

,,  29.     Skull  of  Tritylodon 150 

,,  30.     Molar  of  Tritylodon •         •         -150 

„  31.     Upper  Cheek-Teeth  of  Plesiadapis 155 

,,  32.     Upper  Cheek -Teeth  of  Microcharus 156 


LIST   OF   ILLUSTRATIONS.  xi 

PAGE 

Fig-  33-  Upper  Molars  of  A rctocyon 159 

,,  34.  Upper  Molar  of  Ancodus          .         .         .   •      .         .         .         .  161 

,,  35.  Skeleton  of  Elotherinm 162 

„  36.  Upper  Cheek-Teeth  of  Palaotheriiim 166 

„  37.  Upper  Cheek-Teeth  of  A nchitherium        ,         .         .         .         .  167 

,,  38.  Upper  Molar  of  Horse 167 

„  39.  Skull  of  Hipparion  .         .         .      >.  ,         .         .         ...         .168 

,,  40.  Upper  Molar  of  Rhinoceros       .         .        ..'  ,      i,  .   •„  »•;- .  ,*.,..  169 

,,  41.  Skeleton  of  Titanotherium       .         .         .    •     » .-•   k,        .  171 

,,  42.  Upper  Molar  of  Mastodon         .         .         .. :   .;•...:  •  >.•"••.  -      .  173 

,,  43.  Upper  Cheek-Teeth  of  Coryphodon .         .  •<•    .,...»?!.;*         .  174 

,,  44.  Upper  Cheek-Teeth  of  Anoplotherium      ..  \    ,'.  ,  .  , ,. ... ;       .  185 

,,  45.  Skull  of  PalcBorias    .        •+<  -•;  *,< '.;/•»*-    .1.    .     ».       ..    •'.   ..        .  199 

, ,  46.  Upper  Molar  of  Merycopotanius        .         .         :.     1    •    <    c,    .     .  203 

,,  47.  Restoration  of  Sivatheriutn       .         .         .    .  -.  .      »v  •/»...     .  205 

,,  48.  Skull  of  Lemur         .  ,      »'        .         .         .    .     .;        •       '»    ;     .  217 

,,  49.  Ring-tailed  Lemur  (Lemur  catta)      *  .  ;  «'\  '.• '{.    •     rt&l  .         .  218 

,,  50.  Skull  of  Aye-aye  (Chiromys  madagascariensis}         .«,,..         .  219 

,,  51.  The  Fossa  (Cryptoproctaferooc]         .         ...    *,;,.,    •    ••        •  221 

,,  52.  African  Jumping-Shrew  (Macroscelides  tetradactylus}         .         .  232 

>»  53-  West  African  Potamogale  (Potamogale  velox]         ;.-. ...         .  233 

„  54.  Cape  Hunting-Dog  (Lycaon  pictus]       .  .i   .......     .  236 

,,  55.  Fulgent  African  Flying  Squirrel  {Anomahirus  fiilgens)     .         .  237 

,,  56.  Head  of  African  Wart- Hog  (Phacochcerus  cethiopicus)        .         .  241 

,,  57.  Water-Chevrotain  (Dorcatherium  aquaticuni)  ....  243 

,,  58.  Head  of  Gemsbok  (Oryx gazelld)     .         .         .         .         .         .  246 

,,  59.  Cape  Hy rax  (Procavia  capensis)        ....         .         .  248 

„  60.  Slow  Loris  (Nycticebus  tardigradus}          .....  269 

,,  6r.  Tree-Shrew  (Tupaia  tana) 271 

,,  62.  Indian  Sloth-Bear  (Melursus  ursinus) 275 

,,  63.  Indian  Ratel  (Mellivora  ratel)  . 276 

,,  64.  Japanese  Serow  (Nernorh&dus  crispus] 279 

,,  65.  White-bellied  Pangolin  (Manis  trictispis)          ....  283 

,,  66.  Russian  Desman  (Myogale  moschatd]        .         .                  .         .  320 

,,  67.  Head  of  Spanish  Ibex  (Capra  pyrenaica} 323 

,,  68.  Musk-Deer  (Moschus  moschiferus]    .         .                  .         .         .  325 

,,  69.  Rocky  Mountain  Goat  (Haploceros  montanus]  ....  343 

,,  70.  Musk-Ox  (Ovibos  moschatus)    .         .         .         .         .         i         .  346 

, ,  71.  Musquash  (Fiber  zibethicus) .  362 

,,  72.  Face  of  Geomys  bursarius         .         . .  .    *         .         .         .         .  366 

,,  73.  Face  of  Thomomys  talpoides     .         .         .         .         ...        .  366 

„  74.  Foot  of  Geomys 377 

,,  75.  Head  of  Mule-Deer  (Cariacus  macrotis) 368 

,,  76.  Head  of  Prong-buck  (Antilocapra  americana)  ....  369 


xii  LIST   OF   ILLUSTRATIONS. 


PAGE 

Fig.  77. 

Skeleton  of  Patriofelis  ferox     ..... 

•     372 

,.     78. 

Skeleton  of  Agriochcerus  latifrons     .... 

•     374 

»•     79- 

Hind-foot  of  Agriochcerus         ..... 

•         •     376 

,.     80. 

Skull  of  Protoceras            ...... 

•         •     376 

,,     81. 

Molar  of  Palceosyops         

•     377 

„     82. 

Extremity  of  Skull  of  Uintatherium 

•         •     378 

Figs,  i,  4,  22,  28,  32,  48,  49,  50,  52,  53,  54,  55,  57,  59,  60,  63,  65,  66,  68, 
70,  71  are  from  The  Study  of  Mammals >  Living  and  Extinct,  by  Flower  and 
Lydekker.  Figs.  13,  14,  15,  16,  17,  18,  19,  20,  21,  23,  24  are  taken  from  the 
author's  work  on  Argentine  Fossil  Mammals,  published  in  the  An.  Mus.  La 
Plata.  Figs.  9,  25,  26,  27  are  from  Senor  Ameghino.  Fig.  67  is  from  a  photo- 
graph by  Mr  Abel  Chapman.  Fig.  82  is  from  Prof.  Cope,  fig.  81  from  Prof. 
Earle,  fig.  40  from  Prof.  Boyd  Dawkins,  fig.  30  from  Prof.  Fraas,  and  figs.  7,  8 
from  Prof.  Goodyear.  Fig.  46  is  taken  from  a  plate  in  Hutchinson's  Extinct 
Monsters ;  58  from  a  photograph  by  Mr  Eccles,  of  Reading,  and  75  from  one 
in  the  possession  of  Mr  J.  E.  Harting.  Figs.  6,  35,  41,  43  are  from  Prof. 
Marsh;  and  n,  12,  31,  33,  37,  78,  79,  80  from  Prof.  H.  F.  Osborn.  For  the 
blocks  of  figs.  72,  73,  74,  the  author  is  indebted  to  Dr  C.  H.  Merriam. 
Fig.  45  is  from  Nicholson  and  Lydekker's  Manual  of  Paleontology  ;  fig.  29 
from  Owen;  fig.  64  from  Dr  Sclater  in  the  Proc.  Zool.  Soc.;  fig.  77  from  Dr 
Wortman,  and  39  from  Prof,  von  Zittel. 


A    GEOGRAPHICAL 
HISTORY   OF    MAMMALS. 


CHAPTER  I. 

INTRODUCTORY. 

Distributional  Area  and  Station — Influence  of  Temperature — Humidity — 
Other  Factors  in  Distribution — Importance  of  Palaeontology — Inequality 
in  the  Ages  of  different  Groups  of  Animals — Different  Groups  have  different 
Geographical  Distribution — Importance  of  Mammals  in  Geographical 
Distribution — Classification  of  Mammals — Barriers  to  Dispersal  of  Mam- 
mals— Influence  of  Man  on  Distribution — Extinction  of  the  larger 
Plistocene  Mammals — Distributional  Areas  of  Genera  and  Species — 
Centres  of  Evolution— Permanency  of  Continents  and  Ocean-Basins — 
Zoological  Realms  and  Regions. 

THAT  there  are  differences  in  the  animals  and  plants  of  different 
districts  and  different  countries  is  a  fact  apparent  to  every  person 
who  has  travelled  at  all;  while  to  those  who  have  travelled  ex- 
tensively it  will  further  be  evident  that  the  amount  of  this  difference 
is  by  no  means  correlated  with  the  distance  of  one  country  from 
another,  the  fauna  of  Japan,  for  instance,  being  much  more  like 
that  of  England  and  France  than  is  the  fauna  of  Eastern  Africa  to 
that  of  the  adjacent  island  of  Madagascar.  Unfortunately,  among 
persons  who  are  not  conversant  with  the  principles  of  zoological 
science,  the  distinction  between  the  faunas  of  different  countries 
has  been  much  obscured  by  the  practice  common  to  almost  all 
the  old  voyagers  and  colonists  of  bestowing  upon  the  animals  of 
new  countries  the  names  of  such  Old  World  creatures  as  they 
appeared  most  nearly  to  resemble.  The  puma  of  America  was, 
for  instance,  called  the  lion,  and  the  jaguar  of  the  same  country, 
the  tiger;  while  the  koala  of  Australia  was  christened  the  native 
bear,  and  its  marsupial  allies  the  dasyures  are  still  commonly 
spoken  of  as  native  cats.  To  students  of  the  science  of 
L.  I 


INTRODUCTORY.  [CHAP. 


Geographical  Distribution  it  is,  therefore,  essential  to  discard  such 
misleading  popular  titles,  and  to  speak  of  animals  by  their  correct 
names. 

Apart  from  the  specific   or  generic  distinctions  between  the 
animals  of  one  country  and  another,  the  observer 

Distributional 

Area  and  sta-  will  not  fail  to  notice  more  or  less  well-marked 
differences  between  those  inhabiting  different  dis- 
tricts of  a  single  country ;  such  differences  being  most  intensified 
when  a  country  presents  great  variation  in  its  physical  features. 
An  excellent  instance  of  this  is  afforded  by  South  America,  where 
there  are  the  open  grassy  plains  of  the  Argentine,  the  dense  tropi- 
cal forests  of  Paraguay  and  Brazil,  and  the  snow-clad  heights  of 
the  Andes.  In  the  former  tract  the  traveller  will  meet  with  the 
peculiar  rodents  known  as  viscachas,  the  Patagonian  cavy,  a  species 
of  deer,  numerous  armadillos,  and  the  rhea  (miscalled  the  American 
ostrich).  In  the  Brazilian  forests,  on  the  other  hand,  he  will  find 
monkeys,  marmosets,  tapirs,  tree-porcupines,  sloths,  and  anteaters, 
together  with  certain  armadillos  which  are  for  the  most  part  speci- 
fically or  generically  distinct  from  those  of  the  pampas.  If,  on 
the  other  hand,  he  ascend  high  on  the  Andes,  he  will  leave 
behind  the  animals  of  the  forest,  to  be  confronted  with  chinchillas, 
guanacos  and  vicunas.  Different,  however,  as  are  the  animals  of 
these  various  districts,  yet  an  acquaintance  with  their  zoological 
affinities  will  prove  that  many  of  them  belong  to  closely  allied 
groups,  some  of  which  are  met  with  in  no  other  parts  of  the  world. 
This  will  serve  to  show  that  they  belong  to  what  is  known  as  one 
zoological  province  or  region,  and  that  the  differences  between  the 
faunas  of  different  districts  of  that  province  are  due  to  the  physical 
variations  between  its  component  districts. 

Perhaps  this  point  may  be  still  better  illustrated  by  the  cases 
where  the  same  species  of  animal  is  restricted  to  different  districts 
of  one  country  or  continent.  For  instance,  the  common  squirrel 
is  only  found  in  the  wooded  districts  of  Europe,  and  is  entirely 
absent  from  open  plains.  The  chamois,  again,  is  only  met  with 
in  the  isolated  mountain  ranges  of  the  Pyrenees,  the  Alps,  and  the 
Caucasus  ;  while  the  Siberian  ibex  of  the  Altai  reappears  in  Tibet 
and  the  Himalaya,  but  is  wanting  in  the  intervening  tracts.  In 
these  instances  Europe  would  be  spoken  of  as  the  distributional 


I.]  INFLUENCE   OF   TEMPERATURE.  3 

area  of  the  squirrel  and  the  chamois,  and  Central  Asia  as  that  of 
the  Siberian  ibex ;  but  the  particular  districts  suited  to  the  exist- 
ence of  each  would  be  termed  its  station.  And  here  it  may  be 
mentioned  that  whereas  the  distributional  area  of  a  species  is 
generally  continuous,  its  various  stations  may  be  partially  or  com- 
pletely isolated,  as  in  the  instances  of  the  aforesaid  mountain 
animals,  which  cannot  live  on  the  plains  below. 

Station  is  thus  seen  to  be  very  intimately  connected  with 
temperature;  and  of  this  a  very  striking  example  may 
be  found  among  the  mammals  of  South  America. 
As  already  mentioned,  the  llama-like  animals  respec- 
tively known  as  vicunas  and  guanacos  are  met  with  in  company 
on  the  highlands  of  the  Cordillera  in  Peru  and  Ecuador,  but  as 
we  go  further  south  the  latter  are  found  on  the  plains  of  southern 
Argentina  and  Patagonia,  as  well  as  in  the  island  of  Tierra-del- 
Fuego,  at  the  sea-level.  Here,  then,  there  is  a  clear  proof  of  the 
intimate  connection  existing  between  temperature  and  station; 
the  guanaco,  being  an  animal  which  can  live  only  in  cold  or 
temperate  climates,  finds  suitable  conditions  for  its  existence 
in  tropical  latitudes  solely  at  a  height  of  many  thousands  of  feet, 
although  further  south  it  is  able  to  thrive  at  the  sea-level. 

This  being  so,  it  is  obvious  that  temperature  must  likewise 
exert  a  very  considerable  influence  on  the  whole  distributional 
area  of  many  animals.  Of  this,  the  most  marked  instance  is  found 
in  the  fauna  of  the  Arctic  regions,  which  forms  a  circumpolar  zone 
of  animals  more  or  less  markedly  distinct  from  those  dwelling 
further  south.  And  if  the  whole  land-area  of  the  world  were  con- 
nected, and  not  broken  up  by  mountain-chains,  its  faunas  might 
probably  be  divided  into  zones  or  belts,  whose  limits  would  mainly 
depend  upon  temperature. 

In  this  connection  it  may  be  mentioned  that  the  instance  of  the 
range  of  the  guanaco  is  of  considerable  importance  in  regard  to 
a  decided  difference  between  the  Old  and  New  Worlds  in  respect 
to  the  influence  of  mountains  on  the  present  distribution  of  the 
animals  of  the  two  areas.  In  the  Old  World  the  chief  mountain- 
ranges,  such  as  the  Pyrenees,  Alps,  Carpathians,  Caucasus,  Hindu- 
Koh,  Himalaya,  Thian  Shan,  and  Altai,  run  in  a  more  or  less 
decided  east-and-west  direction;  whereas  in  America,  and  more 

I — 2 


INTRODUCTORY.  [CHAP. 


especially  in  the  southern  half  of  that  continent,  they  have  a  north- 
and- south  trend.  Consequently,  whereas  in  the  former  area  the 
mountain-ranges  have  acted  as  barriers  to  the  dispersal  of  animal 
life,  there  has  been  no  such  obstacle  to  diffusion  in  America,  and 
animals  have  been  able  to  distribute  themselves  according  to  tem- 
perature-conditions. It  is  in  consequence  of  this  physical  feature 
that  a  single  species  of  cold-loving  animal  like  the  guanaco  can 
range  at  the  present  day  from  the  equator  to  latitude  55°  south, 
whereas  in  the  Old  World  the  common  ibex  is  now  restricted  to 
the  isolated  mountain-ranges  of  Europe,  and  the  Siberian  ibex  is 
confined  to  the  systems  of  the  Himalaya  and  the  Altai.  Such 
isolated  stations  could,  of  course,  only  have  been  reached  during 
a  period  when  the  general  temperature  of  the  northern  hemisphere 
was  much  colder  than  it  is  at  present,  and  the  animals  were  thus 
enabled  to  cross  the  lowlands  from  one  chain  to  the  other ;  and 
that  such  a  cold  period  formerly  existed,  there  is  abundant  evidence 
in  the  traces  of  an  extensive  glaciation  found  over  a  large  portion 
of  Europe  and  Asia.  Although,  therefore,  strictly  speaking,  tem- 
perature has  really  had  as  much  effect  in  the  distribution  of 
animals  in  the  Eastern  Hemisphere  as  in  the  Western,  yet,  since 
the  glacial  epoch,  its  influences  have  been  considerably  masked 
by  the  trend  of  the  chief  mountain-ranges,  and  likewise  by  the 
greater  isolation  of  the  different  countries  of  the  former  area  as 
compared  with  the  latter. 

These  essential  differences  thus  render  it  impossible  to  mark 
out  the  Old  World  in  the  zones  of  animal  distribution  which  have 
been  attempted  for  North  America ;  and  they  will  likewise  serve 
largely  to  explain  the  divergence  of  views  on  this  point  which  may 
be  noticed  between  the  writings  of  Drs  Wallace  and  Merriam. 

The  latter  writer1,  whose  conclusions  are  mainly  based  on  the 
evidence  of  North  American  animals  and  plants,  is  of  opinion 
that  in  the  northern  hemisphere  "  animals  and  plants  are  restricted 
in  southward  distribution  by  the  mean  temperature  of  a  brief 
period  covering  the  hottest  part  of  the  year";  and  it  is  added  that 
in  certain  districts  the  mingling  of  essentially  northern  types  with 
those  characteristic  of  a  more  southerly  zone  is  due  to  the  mean 
temperature  of  the  hottest  part  of  the  year  being  sufficiently  low 
1  Appendix,  No.  20. 


I.]  FACTORS   IN    DISTRIBUTION.  5 

for  the  existence  of  the  former,  while  the  total  quantity  of  heat 
suffices  for  the  latter.  In  other  words,  there  is  a  low  summer 
temperature  combined  with  a  high  total  sum  of  heat. 

Among  the  secondary  causes  affecting  distribution,  humidity, 
according  to  the  same  observer,  may  occupy  the  first 
place.  "Humidity,"  he  writes,  " governs  details  of 
distribution  of  numerous  species  of  plants,  reptiles  and  birds,  and 
of  a  few  species  of  mammals,  within  the  several  temperature-zones. 
. . .  Humidity  and  other  secondary  causes  determine  the  presence 
or  absence  of  particular  species  in  particular  localities  within  their 
appropriate  zones,  but  temperature  predetermines  the  possibili- 
ties of  distribution ;  it  fixes  the  limits  beyond  which  species 
cannot  pass ;  it  defines  broad  trans-continental  belts  within 
which  certain  forms  may  thrive  if  other  conditions  permit,  but 
outside  of  which  they  cannot  exist,  be  the  other  conditions  never 
so  favourable." 

Important  as  the  influence  of  temperature  and,  in  a  smaller 
degree,  that  of  humidity,  has  undoubtedly  been  in       other  Fac 
determining  the  distributional  limits  of  the  species     tors  in  Distri- 

f.  ,  ,       ,  .    ,     ,  .  .  bution. 

or  genera  ot  animals  and  plants  now  inhabiting 
particular  countries,  and  large  as  has  been  the  part  played  by  the 
glacial  epoch  in  producing  the  present  condition  of  things,  it  is 
evident  that  temperature  has  been  by  no  means  the  only,  even  if 
it  be  the  chief  factor  in  distribution.  In  the  first  place  there  are 
several  species,  more  especially  among  the  carnivorous  mammals, 
which  seem  quite  independent  of  both  station  and  temperature, 
the  New  World  puma  ranging  from  Patagonia  to  Canada,  while 
the  tiger  inhabits  alike  the  burning  jungles  of  India  and  Burma, 
and  the  Arctic  tundras  of  Siberia.  Striking  as  such  cases  are,  they 
are,  however,  to  be  regarded  merely  as  examples  of  the  individual 
adaptability  of  certain  species,  which,  like  the  carnivores  named, 
are  able  to  obtain  suitable  food  in  any  part  of  the  world,  and 
they  do  not  throw  any  discredit  on  the  power  of  temperature  as 
a  controlling  factor  in  animal  distribution  generally. 

Of  the  utmost  importance  in  this  respect  are  the  changes 
which  the  surface  of  the  globe  itself  has  undergone  in  past 
epochs,  whereby  continents  that  are  now  more  or  less  completely 
sundered  from  one  another  were  formerly  connected,  while  what 


INTRODUCTORY.  [CHAP. 


are  now  islands  were  once  parts  of  continents,  and  vice  versa. 
Such  connections  and  disconnections,  by  allowing  migrations  at 
one  time,  and  preventing  them  from  taking  place  in  the  reverse 
direction  at  a  subsequent  epoch,  have  been  the  chief  factors  which 
have  resulted  in  the  present  very  remarkable  difference  in  the 
faunas  of  different  parts  of  the  globe.  And  it  is  solely  due  to  such 
changes  that  many  of  the  lower  types  of  mammalian  life,  like  the 
marsupials  of  Australia,  and  the  lemurs  and  insectivores  of  Mada- 
gascar, have  been  preserved  at  the  present  day ;  their  insulation 
having  afforded  them  protection  from  the  invasion  of  the  larger 
and  more  specialised  mammals  of  other  parts  of  the  world,  by 
which  they  would  inevitably  have  been  swept  away,  had  the  two 
groups  ever  come  in  contact.  It  is  in  regard  to  these  migratory 
movements  of  animals  and  changes  in  the  land-surface  of  the 
globe  that  zoology  and  geography  are  brought  into  such  close 
relationship ;  the  former  science  sometimes  helping  to  explain  the 
alterations  that  have  taken  place  in  the  contours  of  the  land,  while 
in  other  cases  the  present  distribution  of  the  land  explains  the 
past  history  of  the  animals  by  which  it  is  inhabited. 

To  understand  rightly  the  present  distribution  of  animals,  it  is, 
im  ortance  however,  essential  to  study  their  past  history  as 
of  Paiaeonto-  recorded  by  the  preservation  of  their  fossilised 
remains  in  the  strata  of  the  earth's  crust ;  as  with- 
out such  history  it  would  be  quite  impossible  to  grasp  the  reason 
of  many  apparent  anomalies  in  their  present  distribution.  How, 
for  instance,  without  the  aid  of  palaeontology  would  it  be  possible 
to  understand  how  it  came  about  that  tapirs  are  now  found  only 
in  tropical  America  and  the  Malayan  countries,  or  that  marsupials 
occur  solely  in  America  and  Australasia  at  the  present  day  ?  And 
here  it  may  be  well  to  mention  that  the  science  of  geographical 
distribution  depends  essentially  upon  a  belief  on  the  part  of  the 
student  that  all  animals  are  genetically  connected  one  with 
another,  and  that  the  existing  forms  have  originated  from  earlier 
kinds  by  some  mode  of  evolution.  Were  this  belief  not  accepted, 
the  whole  science  of  distribution  would  fall  to  pieces  ;  as  if  animals 
were  separately  created,  there  would  be  nothing  calling  for  special 
explanation  in  the  fact  of  tapirs  being  restricted  to  the  two  areas 
mentioned. 


I.] 


IMPORTANCE  OF   PALEONTOLOGY. 


To  those  readers  who  may  not  be  geologists,  the  following 
table  of  the  leading  divisions  into  which  the  strata  of  the  earth's 
crust  have  been  divided  will  probably  be  advantageous.  Com- 
mencing with  the  highest  beds,  the  series  will  run  in  descending 
order  as  follows,  viz. : — 

Tertiary.          PLISTOCENE. — Cavern  and  River  Deposits. 

PLIOCENE. — The  "Crags"  of  the  East  Coast. 
MIOCENE. — (Eningen  beds  of  Baden. 
OLIGOCENE. — Gypsum   of  Paris   Basin;    Phos- 
phorites of  Central  France. 
EOCENE. — London  Clay. 
Secondary.        CRETACEOUS. — Chalk,  Upper  Greensand,  Gault, 

Lower  Greensand,  and  Wealden. 
JURASSIC. — Purbeck  beds,  Portland  series,  Kim- 
meridge  Clay,  Coral  Rag,  Oxford 
Clay,     Great     Oolite,     Stonesfield 
Slate,  Inferior  Oolite,  Lias. 
TRIASSIC. — New  Red  Sandstone  of  Cheshire. 
Paleozoic.         PERMIAN. — Red  Marls. 

CARBONIFEROUS. — Coal-Measures    and    Moun- 
tain-Limestone. 
SILURIAN. 
ORDOVICIAN. 
,  «  CAMBRIAN. 


Older  rocks  of  Wales  and  the 
Lake-District. 


Before  attempting  to  draw  any  conclusions  as  to  the  former 
configuration  of  the  surface  of  the  earth  from  the 

,.  .  .,,...,...  Inequality  in 

distribution  of  the  animals  now  inhabiting  its  dif-     the  Ages  of  dif- 


ferent countries,  it  is  essential  to  understand  that 
the  different  classes  into  which  vertebrate  animals 
are  divided  (and  these  only  will  be  taken  into  consideration  in  the 
present  volume)  have  a  very  different  past  history;  the  lower 
groups,  such  as  fishes,  reptiles,  and  amphibians  being  much  older 
types  than  mammals  and  birds,  and  having  attained  their  maximum 
development  at  a  time  when  the  two  latter  formed  but  a  small 
minority  of  the  earth's  population. 

There    is    a    considerable   probability   that    at   least    a   very 
large  proportion  of  the  animals  that  have  populated  the  globe  in 


8  INTRODUCTORY.  [CHAP, 

the  later  geological  epochs  originated  high  up  in  the  northern 
hemisphere,  if  not,  indeed,  in  the  neighbourhood  of  the  pole  itself 
(which  is  known  to  have  enjoyed  a  genial  climate  during  the 
Tertiary  period),  and  that  they  gradually  migrated  southwards  in 
a  series  of  waves,  probably  under  pressure  of  the  development  of 
new  and  higher  types  in  high  latitudes;  and  it  is  to  such  southerly 
migrations  that  the  present  marked  differentiation  of  the  fauna  of 
different  parts  of  the  earth's  surface  is  chiefly  due.  Whether  such 
a  northerly  origin  held  good  for  the  terrestrial  life  of  the  Secondary 
epoch,  there  are  no  means  of  determining ;  but  it  would  appear 
that  the  higher  animals  (which  were  chiefly  reptiles)  of  that  epoch 
were  very  similar  throughout  the  world,  and  that  the  differentiation 
of  faunas  had  scarcely,  if  at  all,  commenced.  Instances  of  this 
are  afforded,  as  noticed  in  the  sequel,  by  the  occurrence  of  an 
identical  genus  of  mammal  (Tritylodori)  in  the  lower  Secondary 
rocks  of  Europe  and  South  Africa ;  as  well  as  by  the  close  alliance 
between  the  dinosaurian  reptiles  from  the  Jurassic  rocks  of 
Europe,  North  America,  Argentina,  India,  and  Madagascar  (the 
genera  being  in  some  cases  identical),  and  likewise  between 
the  anomodont  reptiles  of  the  Trias  of  Europe  and  the  early 
Secondary  rocks  of  South  Africa  and  India. 

Reptiles  belonging  to  orders  still  existing,  such  as  crocodiles 
and  chelonians  (tortoises  and  turtles),  had  already  attained  a  high 
degree  of  development  in  the  Eocene  division  of  the  Tertiary 
period,  when  many  genera  now  living  had  already  made  their 
appearance,  whereas  at  that  time  the  mammals  were  quite  differ- 
ent from  the  modern  forms.  At  the  same  time  the  side-necked 
tortoises  (Pleurodira)  were  the  dominant  forms  in  the  northern 
hemisphere,  whereas  they  have  now  all  migrated  to  southern  lands, 
their  place  in  the  north  being  taken  by  the  more  specialised 
S-necked  group  (Cryptodira).  This,  however,  is  not  all,  for  the 
rhynchocephalians,  of  which  the  sole  existing  representative  is 
the  New  Zealand  tuatera  (Sphenodori),  attained  their  maximum 
development  in  the  northern  hemisphere  during  the  early  part  of 
the  Secondary  epoch,  and  their  southern  migration  must  have 
taken  place  during  some  portion  of  the  same  period.  The 
palaeontological  history  of  amphibians  is  still  very  imperfectly 
known,  but  since  the  group  as  a  whole  is  an  ancient  one,  the 


I.]  DISTRIBUTION    OF   GROUPS   DISSIMILAR.  9 

migrations  of  the  earlier  forms  must  likewise  probably  have  taken 
place  at  an  early  epoch. 

With  mammals  the  case  is  very  different.  The  earliest  known 
forms,  which  date  from  the  Triassic  and  Jurassic  rocks,  are  chiefly 
marsupials  and  forms  apparently  allied  to  the  monotremes,  and  it 
is  probable  that  most  of  the  descendants  of  these,  as  is  more  fully 
indicated  in  the  sequel,  migrated  southwards  during  the  early  part 
of  the  Tertiary  epoch,  to  find  in  Australasia  a  refuge  from  the 
competition  of  higher  forms.  Of  the  higher  placental  mammals, 
none  of  the  modern  types  make  their  appearance  before  the 
Oligocene  and  Miocene  periods,  while  many  do  not  antedate  the 
Pliocene.  Their  southern  migrations  accordingly  took  place  later 
on  in  the  Tertiary  period,  one  of  the  earliest  movements  being  the 
wandering  of  lemuroids,  insectivores,  and  civet-like  carnivores 
into  South  Africa  and  Madagascar.  On  the  other  hand,  many 
other  higher  types,  such  as  the  hippopotami,  giraffes,  and  antelopes, 
which  were  abundant  in  Europe  and  southern  Asia  during  the 
Pliocene,  only  left  their  more  northern  homes  to  find  a  permanent 
abiding  place  in  Africa  at  a  very  late  epoch  in  the  earth's  history. 

Although  the  glacial  epoch  probably  had  a  large  share  in  the 
southern  movements  of  the  later  Tertiary  mammals,  some  cause 
with  which  we  are  unacquainted  would  appear  to  have  been  the 
impelling  power  at  earlier  epochs.  But  be  this  as  it  may,  it  is 
quite  evident  that  a  continuous  series  of  waves  of  migrations  of 
animal  life  has  taken  place  throughout  a  very  long  portion  of  the 
earth's  history.  Similar  migrations  are  also  evident  in  the  case  of 
birds  (which  are  likewise  a  modern  group),  many  forms,  such  as 
secretary-birds  and  trogons,  now  exclusively  southern  in  their 
distribution,  being  represented  in  Europe  during  the  middle  part  of 
the  Tertiary  period. 

From  this  inequality  in  the  ages,  and  consequently  in  the  date 
of  migration,  of  different  groups  of  animals,  it  is 
manifest  that  there  will  be  great  differences  in  the 


present  distribution  of  such  groups;  and  hence  it     different 

.  .      ..  ,       Geographical 

will  be  evident  that  zoological  provinces  indicated     Distribution. 
by  one  group  will  not  hold  good  for  others.  Notable 
instances  of  this  are  afforded  by  the  very  different  divisions  into 
which  the  globe  is  divided  by  those  who  take  mammals,  reptiles, 


10  INTRODUCTORY.  [CHAP. 


or  amphibians  as  their  standards.  Between  birds  and  mammals, 
as  might  have  been  expected  from  the  comparatively  recent  high 
development  of  these  groups,  there  is  much  greater  accord. 
Birds,  however,  differ  from  mammals  (with  the  exception  of  bats) 
in  their  power  of  flight,  which  enables  many  of  them  to  cross 
wide  ocean-tracts,  and  therefore  renders  them  less  valuable  as 
indicative  of  the  changes  that  have  taken  place  in  the  distribution 
of  land  and  water  than  the  latter,  which,  as  a  rule,  require 
direct  means  of  land  transit  for  their  wanderings. 

Excluding  man  (and  for  the  most  part  bats)  and  likewise  the 

aquatic  forms,  such  as  seals,  whales,  and  porpoises, 

o/ Mamma"       niammals  are  the  animals  best  adapted  for  parcelling 

in  Geographi-      out  the  globe  into  zoological  provinces  for  two  chief 

cal  Distribu-  3    _.       .          ,          r  ,  .    ,  , 

tion.  reasons.     Firstly,  they  form  a  group  which  only  at- 

tained its  maximum  development  at  a  comparatively 
late  epoch  of  the  earth's  history;  and,  secondly,  their  movements 
are  mainly  limited  by  the  extent  of  the  land-surfaces  of  the  globe 
which  were  in  actual  communication  at  the  time  of  such  migra- 
tions. Consequently  they  afford  the  safest  and  truest  indications 
of  the  latest  changes  which  have  taken  place  in  the  distribution 
of  land  and  water. 

As  reference  in  the  following  pages  will  constantly  have  to  be 

.fi  made  to  the  various  groups  of  mammals,  it  will  be 

tion  of  Mam-       well  to  give  a  list  in  this  place  of  the  chief  ordinal 

and    subordinal    groups    into   which    the   class   is 

divided ;  such  as  are  now  extinct  being  indicated  by  an  asterisk. 

The  list  stands  as  follows,  viz.  : — 

i.     Order  PRIMATES. — Apes,  Monkeys,  and  Lemurs. 

1  Suborder  ANTHROPOIDEA. — Apes  and  Monkeys. 

2  „         LEMUROIDEA.— Lemurs. 

ii.     Order  CHIROPTERA. — Bats. 

1  Suborder  MEGACHIROPTERA. — Fruit-bats. 

2  ,,         MICROCHIROPTERA. — Insectivorous  Bats. 

iii.     Order  INSECTIVORA. — Insectivores. 

1  Suborder  DERMOPTERA. — Flying-Lemurs. 

2  ,,         INSECTIVORA     VERA. —     Shrews,     Moles, 

Hedgehogs,  etc. 


I.]  CLASSIFICATION    OF   MAMMALS.  II 

iv.     Order  CARNIVORA. — Carnivores. 

1  Suborder  CARNIVORA    VERA. — Cats,    Dogs,    Bears, 

Weasels,  etc. 

2  „         PINNIPEDIA. — Seals  and  Walruses. 

3  „       *CREODONTA. — Hyanodon,  etc. 

v.     Order  RODENTIA. — Rodents. 

1  Suborder  SCIUROMORPHA. — Squirrels,  Marmots,  and 

Beavers. 

2  „          MYOMORPHA. — Dormice,    Mice,    and    Jer- 

boas. 

3  „          HYSTRICOMORPHA. — Porcupines,      Agutis, 

Cavies,  etc. 

4  ,,          LAGOMORPHA. — Picas  and  Hares. 

vi.     Order  UNGULATA. — Hoofed  Mammals. 

1  Suborder  ARTIODACTYLA. — Antelopes,  Camels,  Pigs, 

etc. 

2  ,,         PERISSODACTYLA. — Horses,     Tapirs,     and 

Rhinoceroses. 

3  ,,  *LITOPTERNA. — Macrauchenia,  etc. 

4  „  *ASTRAPOTHERIA. — Astrapotherium,  etc. 

5  „  *PYROTHERIA. — Pyrotherium. 

6  „  *TOXODONTIA. —  Toxodon,  etc. 

7  ,,         HYRACOIDEA. — Hyraces. 

8  ,,         PROBOSCIDEA. — Elephants  and  Mastodons. 

9  ,,  *AMBLYPODA. — Coryphodon,  etc. 

vii.     Order  SIRENIA. — Manatis  and  Dugongs. 
viii.         „      CETACEA. — Whales  and  Porpoises, 
ix.         „      EDENTATA. — Edentates. 
x.         ,,      EFFODIENTIA. — Aard-varks  and  Pangolins. 
xi.         ,,      MARSUPIALIA. — Pouched  Mammals. 

1  Suborder  DIPROTODONTIA. — Kangaroos,  Phalangers, 

and  Wombats. 

2  ,,         POLYPROTODONTIA. — Dasyures,  Bandicoots, 

etc. 

xii.     Order  MONOTREMATA. — Egg-laying  Mammals, 
xiii.         „    *MULTITUBERCULATA. — Pldgiciulax,  Tritylodon,  etc. 


12  INTRODUCTORY.  [CHAP. 


These  orders  are  further  subdivided  into  families  and  genera. 
In  regard  to  the  number  of  the  latter  of  these,  there  is  still  a  con- 
siderable difference  of  opinion  among  naturalists  ;  but  in  the 
present  work  those  adopted  in  Flower  and  Lydekker's  Study  of 
Mammals1  will  be  in  the  main  adhered  to,  with  such  corrections 
and  additions  as  are  necessary  owing  to  recent  emendations  in 
nomenclature  or  to  the  discovery  of  new  forms. 

Omitting  from   consideration   the  purely  aquatic  and  volant 

members  of  the  class,  the  most  effectual  barriers  to 

Dispersal  of        the  dispersal  of  mammals  are  formed  by  channels 

Mammals. 


their  being  crossed  by  swimming.  And  it  is  this  inability  to 
traverse  any  extent  of  water  that  renders  what  are  known  as 
oceanic  islands  practically  devoid  of  all  mammalian  life,  with 
the  exception  of  a  few  bats  and  small  rodents  ;  the  latter  animals 
having  apparently  some  means  of  dispersal  not  common  to  other 
members  of  this  class.  Oceanic  islands,  it  may  be  explained,  are 
such  as  rise  from  great  depths  in  the  ocean,  and  are  composed, 
almost  invariably,  either  of  volcanic  rocks  or  of  coral.  They  show, 
for  the  most  part,  no  decisive  evidence  of  having  been  connected 
with  any  continental  land,  and  thus  have  never  been  enabled  to 
receive  a  mammalian  fauna2.  In  marked  distinction  to  these  are 
the  so-called  continental  islands,  such  as  Madagascar  and  Great 
Britain,  which,  both  from  the  evidence  of  their  mammalian  fauna 
and  their  geological  conformation,  have  indubitably  been  in  direct 
communication  with  the  adjacent  continent  at  no  very  distant 
epoch.  As  a  rule,  the  channels  between  such  islands  and  the 
mainland  are  comparatively  shallow,  so  that  a  moderate  degree  of 
upheaval  would  place  the  two  in  direct  connection. 

The  relative  depth  of  the  channel  between  two  islands,  or 
between  an  island  and  a  continent  is  indeed  of  much  more  im- 
portance in  regard  to  the  dispersal  of  mammals  than  is  its  width. 
This  is  best  exemplified  by  the  well-known  case  of  "Wallace's 
line  "  in  the  Malayan  archipelago  ;  that  name  being  applied  to 

1  London,  1891. 

2  There  is  a  possibility  that  some  oceanic  islands  may  have  been  connected 
with  continents,  and  that  their  original  mammalian  fauna  has  been  destroyed 
by  submergence. 


I.]  BARRIERS   TO   DISPERSAL.  13 

the  narrow  strait  separating  the  islands  of  Bali  and  Lombok,  and 
its  northward  continuation,  the  Makassar  Strait,  dividing  Borneo 
from  Celebes.  Although  the  two  former  islands  are  extremely 
close  together,  while  Celebes  is  much  less  widely  separated  from 
Borneo  than  is  the  latter  from  Sumatra,  yet  the  faunas  of  Lombok 
and  Celebes  are  markedly  distinct  from  those  of  the  islands  lying 
to  the  north  and  west  of  Wallace's  line.  Soundings  show  that  the 
Makassar  Strait,  and  likewise  the  Bali-Lombok  Strait  are  of  greater 
depth  than  the  channels  separating  the  other  islands  of  the  archi- 
pelago; and  consequently  that  Wallace's  line  indicates  a  very 
old  barrier  which  has  long  been  impassable  to  the  majority  of 
mammals. 

That  continental  islands  have  received  the  great  bulk  of  their 
mammalian  fauna  by  means  of  a  more  or  less  complete  land- 
connection  with  the  mainland,  is  perfectly  evident.  Nevertheless, 
there  are  cases  where  certain  mammals  have  crossed  the  inter- 
vening channel,  either  by  swimming,  or  by  having  been  carried 
across  on  natural  rafts  of  some  kind ;  an  instance  of  this  nature 
being  exemplified  by  the  occurrence  of  an  African  type  of  pig  in 
Madagascar.  It  thus  becomes  a  question  of  considerable  interest 
to  ascertain  what  stretches  of  sea  large  mammals  are  capable  of 
crossing.  It  is  stated  that  the  jaguar  has  been  known  to  swim 
across  the  Rio  de  la  Plata,  which  at  its  mouth  is  something  like 
eighty  miles  across ;  and  a  polar  bear  has  been  observed  swim- 
ming at  a  distance  of  twenty  miles  from  land  in  Bering  Strait. 
The  tiger  frequently  crosses  the  narrower  channels  in  the  Sandar- 
bans  of  Lower  Bengal;  and  both  deer,  pigs,  and  elephants  are 
good  swimmers.  The  latter  animals  have,  indeed,  been  known  to 
swim  for  six  hours  at  a  stretch,  and,  with  a  rest,  for  upwards  of 
nine ;  but  their  rate  of  progress  is  extremely  slow,  and  probably 
exceeds  but  little,  if  at  all,  a  mile  an  hour.  The  Palk  Strait, 
which  is  considerably  less  than  forty  miles  in  width  at  its  narrowest 
part,  has  formed  an  effectual  barrier  to  the  passage  of  the  tiger 
from  India  into  Ceylon;  and  it  may  accordingly  be  assumed 
that  about  twenty  miles  is  the  utmost  limit  which  mammals  are 
likely  to  cross  by  swimming,  even  when  favoured  by  currents. 

Such  passages  as  these  must,  however,  be  of  very  rare  occur- 
rence, for  a  terrestrial  mammal  is  not  likely  to  take  it  into  its  head 


14  INTRODUCTORY.  [CHAP. 

to  swim  straight  out  to  sea  in  an  unknown  direction.  Moreover, 
supposing  a  mammal  new  to  a  particular  island  to  have  arrived 
there  by  swimming,  unless  it  happen  to  be  a  pregnant  female,  or 
unless  another  individual  of  the  same  species  but  of  the  opposite 
sex  should  arrive  soon  after  (a  most  unlikely  event),  it  would  in 
due  course  die  without  being  able  to  propagate  its  kind.  And 
even  if  it  should  happen  to  be  a  pregnant  female,  there  would  be 
no  certainty  that  its  offspring,  if  but  one  in  number,  should  be  of 
the  opposite  sex  to  its  parent.  Accordingly,  it  would  seem  that 
the  population  of  islands  by  mammals  that  have  arrived  by  swim- 
ming must  be  a  very  rare  event  indeed.  Rafts  may  be  of  more 
importance.  Mr  Aplin,  in  the  Proc.  Zool.  Soc.  for  1894,  mentions 
that  jaguars  and  pumas  are  frequently  transported  by  them  from 
one  side  of  the  Rio  de  la  Plata  to  the  other;  and  in  the  rainy 
season  many  are  to  be  seen  off  the  northern  coast  of  Borneo, 
some  of  which  may  be  as  much  as  thirty  yards  in  length.  Still 
such  rafts  are  not  likely  to  cross  straits,  except  when  there  is  a 
current  setting  from  one  bank  to  the  other. 

Before  leaving  this  part  of  the  subject,  it  must  be  mentioned 
that  the  degree  of  difference  between  the  fauna  of  an  island  and 
that  of  the  adjacent  continent,  or  between  the  faunas  of  two 
islands,  affords  a  most  important  clue  as  to  the  relative  date  of  the 
land-connection  between  them.  Madagascar,  for  instance,  has  a 
mammalian  fauna  which  although  clearly  derived  from  Africa,  is 
yet  so  different  from  that  now  inhabiting  the  mainland,  that  the 
land-connection  between  the  two  must  have  been  broken  up  at  an 
epoch  comparatively  remote.  Ceylon  and  India  present  a  condition 
in  respect  of  their  faunas  intermediate  between  the  last  and  that  of 
Great  Britain  as  compared  to  the  Continent.  In  the  latter  instance, 
with  the  exception  of  a  single  Irish  weasel,  all  the  mammals  are 
identical  with  species  now  inhabiting  the  Continent,  thus  proving 
that  the  connection  has  been  a  comparatively  recent  one ;  although 
the  peculiar  weasel  indicates  that  the  separation  between  Ireland 
and  Britain  has  been  of  sufficient  duration  to  admit  of  the  develop- 
ment of  a  distinct  specific  type  in  the  former  country. 

Although  large  rivers  like  the  Amazon  and  La  Plata  un- 
doubtedly form  serious  barriers  to  the  migration  of  mammals,  yet 
these  are  not  so  insuperable  as  might  at  first  sight  be  supposed, 


I.]  BARRIERS   TO   DISPERSAL.  15 

owing  to  the  fact  that  in  districts  where  vegetation  is  luxuriant, 
huge  natural  rafts  are  formed  by  the  trunks  of  trees  intermingled 
with  vegetable  matter,  upon  which  numbers  of  animals  may  be 
borne  down  stream,  and  thus  transferred  from  one  bank  to  the 
other.  Nevertheless,  in  treeless  districts,  or  near  their  mouths, 
large  rivers  afford  absolutely  impassable  barriers  to  the  movements 
of  mammals.  In  South  America,  for  instance,  even  such  an 
aquatic  creature  as  the  carpincho,  or  capibara  (ffydrochcerus)  has 
been  unable  to  cross  the  Plata  river  from  Uruguay  into  the 
Argentine,  while,  conversely  the  viscacha  (Lagostomus)  of  Argentina 
is  prevented  by  the  same  river  from  reaching  Uruguay. 

Deserts  are,  perhaps,  even  more  impracticable  than  rivers ;  the 
Sahara — which  was  long  supposed  to  have  been  the  site  of  an 
ancient  sea,  although  it  has  really  been  a  desert  since  very  remote 
ages — having  apparently  formed  a  barrier  preventing  the  fusion  of 
the  mammals  of  North  Africa  with  those  to  the  south  of  that  tract 
since  at  least  the  Pliocene  epoch.  It  must  not,  however,  be  sup- 
posed that  what  are  desert-tracts  at  the  present  day  have  always 
been  such,  the  existence  of  a  fossil  chimpanzee  in  North-Western 
India  during  the  Pliocene  period  indicating  that  the  open  sandy 
plains  of  the  Punjab  were  at  that  time  covered  with  dense  tropical 
forests,  and  probably  that  the  same  was  the  case  with  parts  of 
Syria  and  Arabia. 

Before  taking  leave  of  seas  and  deserts,  it  should  be  mentioned 
that  in  the  polar  regions  ice  may  act  in  lieu  of  a  land-connection 
to  enable  mammals  to  pass  from  one  country  to  another.  On  this 
subject  Dr  Heilprin  writes  that  "  the  reindeer  is  stated  to  cross 
the  Bering  Strait  by  way  of  the  Aleutian  Islands  and  the  Frozen 
Sea,  and  in  a  somewhat  similar  manner  the  musk-ox  finds  its  way 
to  Melville  Island;  it  is,  however,  somewhat  singular  that  the  last- 
named  animal,  despite  its  long  ice-journeys,  never  manages  to 
reach  either  the  continent  of  Asia  or  Greenland." 

High  mountain  ranges  form  an  effectual  barrier  to  the  migration 
of  mammals,  not  only  on  account  of  the  physical  difficulties  of 
crossing  them,  but  likewise  by  the  lowness  of  the  temperature  at 
great  altitudes,  coupled  with  the  absence  of  proper  food,  being 
fatal  to  the  existence  of  many.  As  already  stated,  however, 
mountain-ranges  are  much  more  far-reaching  in  their  effects  on  such 


l6  INTRODUCTORY.  [CHAP. 

migrations  when,  as  in  the  Old  World,  their  trend  is  from  west  to 
east,  than  when,  as  in  America,  they  run  from  north  to  south,  and 
thus  do  not  interfere  with  the  free  movements  of  plain-dwelling 
animals  on  either  side.  In  many  instances  the  mammals  inhabit- 
ing each  of  the  isolated  mountain-chains,  as  those  of  Europe,  are 
to  a  great  extent  specifically  identical  one  with  another,  not  having 
had  time  to  become  modified  into  distinct  species  since  they 
reached  their  present  haunts  at  the  close  of  the  glacial  period. 
Even  among  these,  however,  there  are  indications  of  the  com- 
mencement of  specific  differences,  the  chamois  of  the  Caucasus 
forming  a  variety  differing  somewhat  from  the  typical  Alpine  race. 
Where  the  isolation  has  been  longer,  as  in  the  case  of  the  fauna 
of  the  highlands  of  Tibet,  the  difference  is  much  more  strongly 
marked ;  the  mammalian  fauna  in  this  instance  being  as  peculiar 
and  distinct  as  that  of  many  ancient  continental  islands. 

Probably  ever  since  man  has  existed  in  any  numbers  on  the 
influence  of  gl°be  he  has  been  exerting  a  more  or  less  strongly- 
ManonDistri-  marked  influence  on  the  distribution  of  animals, 
either  by  destroying  them,  or  by  conveying  them  to 
countries  or  districts  which  are  not  their  natural  home.  By  the 
involuntary  aid  of  man  the  common  rat  and  mouse,  which  belong 
to  a  genus  unknown  in  the  New  World,  have  been  conveyed  to 
every  country  in  the  globe ;  while  the  rabbit  has  been  carried  to 
the  Antipodes,  where  it  has  flourished  and  increased  in  an 
unprecedented  manner.  Cattle  and  horses  have  been  introduced 
into  South  America,  Australia,  and  other  countries  where  they 
were  naturally  unknown,  and  by  their  rapid  increase  have  shown 
that  the  absence  of  particular  animals  from  particular  districts  is 
not  necessarily  due  to  their  being  unsuited  to  live  there,  but  rather 
to  the  fact  that  they  have  been  unable  to  find  their  way  thither. 
The  fallow-deer,  again,  has  been  imported  from  its  Mediterranean 
home  into  England  and  other  countries  of  northern  Europe;  while 
goats  and  pigs  have  been  carried  to  a  number  of  oceanic  islands, 
where  they  have  done  irreparable  harm  in  exterminating  the  native 
fauna  and  flora. 

In  all  these  instances  the  fact  of  the  introduction  has  always 
been  more  or  less  clearly  known,  and  therefore  no  difficulty  arises 
as  to  what  are  native  and  what  are  introduced  forms.  Very 


I.]  MAN    AND   DISTRIBUTION.  1 7 

different,  however,  is  the  case  with  the  islands  of  the  Malay  Archi- 
pelago, where  the  natives,  who  have  a  wonderful  facility  for 
taming  animals,  have  carried  a  species  peculiar  to  one  district  or 
island  to  localities  where  it  is  quite  unknown  as  a  native ;  and  in 
consequence  of  this  transportation  and  acclimatisation  it  is  pro- 
bable that  several  mammals  have  been  given  a  habitat  to  which 
they  have  not  the  most  remote  right.  To  the  Malays  is  due  the 
introduction  of  the  small  civet  known  as  the  rasse  into  Mada- 
gascar. Whether  the  dingo,  or  native  dog  of  Australia,  was  intro- 
duced at  an  exceedingly  remote  era  by  the  original  colonisers  of 
that  island,  or  whether  it  is  truly  indigenous,  is  a  question  that 
will  probably  never  be  decisively  answered.  It  is  likewise  quite 
impossible  to  say  what  part  man  may  have  played  in  the  extermi- 
nation of  the  large  mammals  that  inhabited  Europe  about  the 
close  of  the  glacial  period,  but  it  seems  quite  probable  that  he 
may  have  had  a  considerable  share  in  their  destruction.  Be  this 
as  it  may,  the  domestication  of  certain  mammals  has  undoubtedly 
had  the  effect  of  destroying  the  wild  race,  as  is  remarkably  ex- 
emplified by  the  two  existing  species  of  camel,  of  neither  of  which 
do  we  know  the  original  habitat.  The  original  European  wild  ox 
—unless,  indeed,  the  half-wild  cattle  of  the  British  parks  be  its 
direct  descendants — has  likewise  disappeared  at  some  unknown 
epoch  owing  to  the  hand  of  man.  Although  other  mammals,  such 
as  the  quagga  (Equus  quagga},  Burchell's  rhinoceros  (Rhinoceros 
simus),  and  the  blaubok  (Hippotragus  leucophaus)  have  been 
almost  or  completely  exterminated  by  human  agency  in  South 
Africa,  while  the  American  bison  has  been  practically  swept  away 
from  its  native  prairies,  yet  in  all  these  instances  there  is  a  more 
or  less  full  record  of  the  original  range  of  the  creatures.  In 
other  cases  also  mammals  have  been  utterly  exterminated  by 
human  agency  from  countries  of  which  they  were  originally  in- 
habitants, as  is  exemplified  by  the  disappearance  from  the  British 
Islands  of  the  bear,  the  wolf,  the  beaver,  and  the  wild  boar  within 
the  historic  period,  although  they  still  survive  in  other  parts  of 
their  habitat.  In  these  particular  instances  there  is  fortunately 
full  evidence  as  to  the  former  existence  of  these  animals  in 
Britain  ;  but  it  is  highly  probable  that  in  more  remote  countries 
mammals  have  been  exterminated  without  any  record  being  left 
L.  2 


1  8  INTRODUCTORY.  [CHAP. 

of  their  existence,  so  that  the  full  extent  of  their  range,  if  they  be 
surviving  forms,  can  now  never  be  ascertained. 

To  quote  the  words  of  Dr  Wallace,  it  is  evident  that  in  the 
present  day  we  live  in  an  impoverished  epoch,  so 

Extinction 

of  the  larger         iar  as  the  larger  mammals  are  concerned,  as  com- 


pared  with  the  Plistocene  era  ;  this  being  true  not 
only  as  regards  the  northern  half  of  the  Old  World, 
but  likewise  North  and  South  America,  as  well  as  Australia. 
From  the  northern  half  of  the  Old  World  have  disappeared  the 
mammoth,  the  elasmothere,  the  woolly  and  other  species  of 
rhinoceros,  the  sabre-toothed  tigers,  etc.;  North  America  has  lost 
the  megalonyx  and  the  Ohio  mastodon  ;  from  South  America  the 
glyptodonts,  mylodons,  the  megalothere,  and  the  macrauchenia  have 
been  swept  away  ;  while  Australia  no  longer  possesses  the  dipro- 
todon  and  various  gigantic  species  of  kangaroos  and  wombats. 
In  the  northern  hemisphere  this  impoverishment  of  the  fauna  has 
been  very  generally  attributed  to  the  effects  of  the  glacial  period, 
but  although  this  may  have  been  a  partial  cause,  it  can  hardly  be 
the  only  one.  The  mammoth,  for  instance,  certainly  lived  during 
a  considerable  portion  of  the  glacial  epoch,  and  if  it  survived  thus 
far,  why  should  it  have  disappeared  at  the  close?  Moreover,  all 
the  European  mastodons  and  the  southern  elephant  (Elephas 
meridionalis]  died  out  before  the  incoming  of  glacial  conditions  ; 
and  the  same  is  true  of  all  the  extinct  elephants  and  mastodons  of 
southern  Asia.  Further,  a  large  number  of  English  geologists 
believe  the  brick-earths  of  the  Thames  valley,  which  contain 
remains  of  rhinoceroses  and  elephants  in  abundance,  to  be  of 
post-glacial  age.  As  regards  the  southern  hemisphere,  it  can 
hardly  be  contended  that  glacial  conditions  prevailed  there  at  the 
same  time  as  in  the  northern  half  of  the  world. 

It  is  thus  evident  that  although  a  very  great  number  of  large 
mammals  were  exterminated  (perhaps  partly  by  the  aid  of  human 
agency)  at  the  close  of  the  Plistocene  period,  when  the  group 
had  attained  its  maximum  development  as  regards  the  bodily  size 
of  its  members,  yet  other  large  forms  had  been  steadily  dying 
out  in  previous  epochs.  And  it  would  seem  that  there  must 
be  some  general  deep-seated  cause  affecting  the  life  of  a  species 
with  which  we  are  at  present'  unacquainted.  Indeed,  as  there 


I.]  DISTRIBUTIONAL  AREAS.  19 


is  a  term  to  the  life  of  an  individual,  what  is  more  natural  than 
that  there  should  also  be  one  to  the  existence  of  a  species  ?  It  still 
remains,  indeed,  to  account  for  the  fact  that  the  larger  Plistocene 
mammals  had  no  successors  in  the  greater  part  of  the  world,  but 
perhaps  this  is  in  some  way  connected  with  the  advent  of  man. 

Before  coming  to  the  consideration  of  the  zoological  divisions 
into  which,  from  the  present  geographical  distribu- 
tion of  mammals,  the  world  may  be  mapped  out,  it     tionai  Areas  of 
is  necessary  to  devote  a  brief  space  to  the  considera-     Genera^ and 
tion  of  two  other  points  ;  the  first  relating  to  the 
relative  size  of  the  distributional  area  of  genera  and  species,  and 
the  second  to  the  permanency  of  ocean-basins  and  continents. 

As  regards  the  first  point,  it  appears  to  be  true  in  the  case  of 
mammals  (although  not  of  all  other  groups)  that  every  species  has 
a  continuous  distributional  area,  except  where  this  has  been  broken 
up  by  human  destructiveness.  It  is  not  meant  by  this  that  every 
part  of  such  area  is  inhabited  by  the  particular  species,  as 
"station"  renders  this  impracticable;  but  merely  that  the  whole 
area  is  ranged  over  by  the  species  in  such  spots  as  are  suited  to 
its  particular  mode  of  life.  Great  variation  obtains,  however,  in 
regard  to  the  size  of  such  distributional  area;  and  it  will  be 
obvious  that  the  size  of  the  area  varies  directly  as  the  adaptability 
of  the  species  to  different  climatic  and  other  physical  conditions. 
Perhaps  the  most  important  condition  of  all  is  the  possibility  of 
obtaining  suitable  food ;  and  in  this  respect  carnivorous  mammals 
are  in  a  far  better  position  than  any  other  members  of  their  class, 
since  the  kind  of  animal  on  which  they  prey  is  immaterial.  This 
will  readily  account  for  the  extensive  geographical  ranges  enjoyed 
by  the  puma  and  the  tiger,  which,  as  stated  on  page  5,  embrace 
almost  every  degree  of  latitude.  Animals  with  such  a  wide  dis- 
tribution are  of  but  little  use  to  the  student  of  geographical 
distribution.  Moreover,  it  will  generally  be  found  that  species 
with  a  wide  range  belong  to  large  genera  having  a  still  more 
extensive  distributional  area;  this  being  markedly  the  case  with 
the  puma  and  the  tiger ;  the  genus  Felts  being  one  of  the  largest 
in  the  class,  and  ranging  over  the  whole  world  with  the  exception 
of  Australasia.  Such  cosmopolitan  genera  are  likewise  almost 
valueless  to  the  distributionist. 

2 — 2 


20  INTRODUCTORY.  [CHAP. 

On  the  other  hand,  species  with  a  small  distributional  area 
usually  belong  to  small  genera,  of  which  they  may  be  the  only 
representatives.  Instances  of  this  nature  are  afforded  by  the 
panda  (sElurus]  and  binturong  (Arctictis)  of  the  eastern  Himalaya, 
and  the  parti-coloured  bear  (^Eluropus]  and  chiru  antelope  (Pan- 
tholops]  of  the  Tibetan  plateau.  Although  such  single  representa- 
tives of  genera  are  highly  important  to  the  study  of  distributional 
zoology,  of  vastly  greater  importance  are  small  genera  having  two 
or  more  species  living  in  widely  separated  areas.  Examples  of 
such  are  to  be  found  among  the  porcupines  of  the  genus  Atherura, 
of  which  one  species  is  Malayan  and  the  other  two  West  and 
Central  African ;  in  the  mice  of  the  genus  Golunda,  with  one 
African  and  one  Indian  representative ;  and  likewise  by  the  tapirs 
(Tapirus\  of  which  there  is  one  Malayan,  and  several  tropical 
American  species.  These  examples  of  "  discontinuous  distribu- 
tion "  among  genera  are  of  the  very  highest  import  to  the  science ; 
since  they  clearly  indicate  that  some  of  the  lands  lying  between  its 
present  disconnected  distributional  areas  must  have  formerly  been 
the  habitat  of  the  genus,  and  thus  enable  important  conclusions  to 
be  drawn  as  to  the  former  land-connections  between  such  areas. 
Both  in  the  case  of  the  tapirs  and  of  the  brush-tailed  porcupines, 
remains  of  extinct  species  have  been  discovered  in  the  intermediate 
areas. 

Equally  important  are  families,  either  large  or  small,  which 
contain  two  or  more  closely  allied  small  genera  respectively  con- 
fined to  distant  areas.  As  an  instance  of  a  large  family  containing 
such  allied  genera,  may  be  cited  the  Viverridce,  among  which  the 
true  linsangs  (Linsangd)  are  represented  by  several  species  from 
the  Eastern  Himalaya  and  the  Malayan  countries,  while  the 
closely-allied  Poianais  confined  to  West  Africa.  The  chevrotains 
(Tragulidee),  on  the  other  hand,  form  a  small  family  with  a  dis- 
continuous distribution  ;  one  genus  (Tragulus)  being  now  Oriental, 
while  the  other  (Dorcatheriuni)  is  West  African.  Here  it  is  quite 
evident,  of  course,  that  the  distributional  area  of  the  family  must 
once  have  been  continuous  ;  and,  as  a  matter  of  fact,  remains  of 
both  genera  occur  in  the  Pliocene  of  India,  those  of  the  latter  being 
also  found  in  the  European  Miocene. 

In  other  families  with  a  discontinuous  distribution,  as  in  the 


I.]  CENTRES    OF    EVOLUTION.  21 

rodent  family  Octodontida,  which  is  now  mainly  confined  to  Africa 
south  of  the  Sahara,  and  Central  and  South  America,  the  genera 
may  be  less  closely  allied,  although  sufficiently  so  to  indicate 
a  continuous  distributional  area,  or  rather  a  common  centre  of 
dispersal,  at  no  very  remote  epoch. 

Allied  families,  with  a  small  number  of  genera,  severally  con- 
fined to  distant  localities  are  likewise  of  the  highest  value  in 
building  up  the  former  history  of  the  globe.  As  examples  of  this 
nature  may  be  cited  the  tree-shrews  (Tupaiida)  of  the  Oriental 
countries  and  the  jumping  shrews  (Macroscelidida)  of  Africa  on  the 
one  hand,  and  the  Solenodontidce  of  the  West  India  islands,  and 
the  Centetidce  of  Madagascar  on  the  other.  Such  families  must 
clearly  have  had  a  common  centre  of  origin  and  dispersal  ;  the 
available  evidence  suggesting  that  in  the  case  of  the  two  former 
such  centre  was  Europe. 

Although  of  far  less  common  occurrence  than  among  families 
or  genera,  discontinuous  distribution  in  an  order  is  perhaps  of  even 
more  importance  than  either  of  the  other  cases,  as  it  implies  a 
greater  interval  of  time  since  the  original  dispersal  took  place,  and, 
therefore,  carries  back  such  conclusions  as  can  be  drawn  in  regard 
to  former  land-connections  to  a  still  earlier  epoch.  Among 
mammals  the  only  instance  of  this  nature  is  to  be  found  in  the 
marsupials1,  of  which  two  families  are  American  (and  mainly 
South  and  Central  American),  while  all  the  others  are  confined  to 
Australasia  and  some  of  the  adjacent  Malayan  islands.  In  this 
case  also  there  is  abundant  evidence  of  the  wide  distribution  of 
the  whole  group  in  former  epochs  of  the  earth's  history. 

This  last  instance  leads  on  to  the  consideration  of  what  may 
be  termed  "centres  of  evolution."  In  a  previous 


paragraph  it  has  been  stated  that,  according  to  the  "  °f 


available  evidence,  a  very  large  proportion,  if  not 
the   whole,   of  the  terrestrial   mammalian  life    of  the  globe  has 
originated  in  the  northern  hemisphere,  from  which  it  has  spread 
southwards  in  a  continuous  successive  series  of  waves.     When, 
however,  certain  groups  of  mammals  had  once  reached  the  more 

1  In  this  work  the  Effodientia  are  separated  from  the  Edentata;  but  when 
these  are  united,  there  is  a  second  instance. 


22  INTRODUCTORY.  [CHAP. 


remote  parts  of  the  southern  hemisphere,  where  they  were  free 
from  the  competition  of  the  higher  forms,  and  met  with  favourable 
conditions,  they  seemed  to  take  a  new  lease  of  life,  and  attained  a 
fulness  and  variety  of  development  which  they  had  never  reached 
before.  As  a  rule,  more  or  less  complete  isolation  has  been  a 
dominant  feature  of  this  development;  of  which  the  best  and  most 
striking  instance  is  that  of  the  marsupials  in  Australasia.  That 
area  may  accordingly  be  called  the  marsupial  evolutionary  centre. 
Scarcely  less  striking  is  the  instance  of  the  edentates  (of  which  the 
original  derivation  is  unknown)  in  South  America,  where,  in 
company  with  certain  peculiar  extinct  groups  of  ungulates,  they 
attained  an  extraordinary  development,  both  as  regards  the 
number  of  specific,  generic,  and  family  types,  and  likewise  in 
respect  of  the  bodily  size  of  some  of  its  members.  This  second 
area  may  be  termed  the  evolutionary  centre  of  the  edentates.  A 
third  great  centre  is  constituted  by  Europe,  Asia,  and  North 
America,  which  appear  to  have  been  the  main  developmental 
centre  of  the  higher  mammals,  and  may  accordingly  be  named  the 
placental  evolutionary  centre.  Two  other  minor  centres  are 
respectively  indicated  by  Madagascar  and  Africa  south  of  the 
Sahara :  the  former  as  being  the  headquarters  of  the  lemurs,  may 
appropriately  be  spoken  of  as  the  lemuroid  centre,  while  the 
great  development  of  the  antelopes  in  Africa  suggests  the  name  of 
the  antelopine  evolutionary  centre  for  that  continent. 

The  circumstance  that  throughout  the  greater  part  of  North 

America  and  Europe  a  very  large  proportion  of  the 

of  Conine ntsy     continents   are   built   up   of  sedimentary  strata  of 

and  Ocean-          marine  origin,  naturally  led  geologists  in  the  early 

Ba.si.ns.  .      .  .  1  1*1 

days  of  their  science  to  the  conclusion  that  every 
part  of  the  land  had  at  one  time  been  deep  ocean,  and  every 
stretch  of  ocean  dry  land.  More  careful  study  led,  however, 
to  the  belief  that  this  idea  was  not  founded  on  fact,  and  that 
although  it  was  perfectly  true  that  what  are  now  continents  had 
been  many  times  under  the  sea,  yet  that  such  areas  had  never 
formed  abyssal  ocean-depths ;  and,  conversely,  that  such  ocean- 
depths  had  never  been  dry  land.  In  addition  to  many  other  lines 
of  evidence,  this  view  of  the  permanency  of  continents  and  ocean- 
basins  is  strongly  supported  by  the  circumstance  that  nearly  all 


I.]  CONTINENTS   AND   OCEAN-BASINS.  23 

oceanic  islands  are  either  of  volcanic  or  coral  origin,  and  do  not 
contain  sedimentary  rocks;  and  also  that  deposits  analogous  to 
those  laid  down  in  the  deepest  ocean  beds  are  generally  wanting 
from  among  the  sedimentary  series  of  rocks  of  which  the  continents 
and  islands  are  composed.  A  further  argument  was  afforded  by 
the  discovery  that  the  greater  portion  of  peninsular  India  and 
South  Africa  has  been  dry  land  since  the  Palaeozoic  epoch. 

As  is  so  commonly  the  case  in  similar  instances,  the  promulga- 
tors  of  the  doctrine  of  the  permanency  of  continents  and  ocean- 
basins  pressed  their  hypothesis  too  far;  and  it  is  now  evident 
that  although  the  doctrine  is  true  as  a  whole,  and  more  especially 
as  regards  the  later  stages  of  the  earth's  history,  yet  it  requires 
very  considerable  modification  from  the  original  form  in  which  it 
was  advanced.  In  the  first  place,  it  has  been  shown  that  crystal- 
line granitic  and  gneissic  rocks  occur  in  the  Seychelles,  which  were 
formerly  regarded  as  true  oceanic  islands ;  and,  secondly,  deep-sea 
deposits  have  been  discovered  in  the  West  Indies  and  the  Solomon 
Islands.  Moreover,  various  lines  of  evidence  indicate  that  during 
the  Jurassic  and  Cretaceous  epochs  there  was  a  continuous  land- 
connection  between  Africa  (by  way  of  Madagascar  and  the 
Seychelles)  and  India;  while  at  some  time  in  the  Secondary  era, 
in  the  opinion  of  Drs  Neumayr  and  Blanford,  South  America  and 
South  Africa  were  in  communication  across  the  South  Atlantic. 
The  latter  connection  appears,  indeed,  to  have  been  a  survival 
from  an  older  Palaeozoic  girdle  of  land  which,  from  the  evidence  of 
fossil  floras,  seems  to  have  existed  in  low  latitudes  round  nearly 
three-quarters  the  circumference  of  the  globe,  and  which  was  cut 
oft"  from  the  land  to  the  north.  There  is,  moreover,  the  possibility 
of  a  Tertiary  connection  of  Australia  with  Patagonia  by  way  of 
Polynesia,  to  which  allusion  is  made  in  the  third  chapter.  Then, 
again,  the  recent  investigations  of  Dr  J.  W.  Gregory1  on  the  fossil 
corals  of  the  West  Indies  have  afforded  strong  support  to  the  view 
that  the  Atlantic  is  of  comparatively  recent  origin.  After  referring 
to  the  remarkable  resemblance  between  the  existing  fauna  of  the 
West  Indian  seas  and  that  of  the  Miocene  deposits  of  the  Mediter- 
ranean basin,  Dr  Gregory2  observes  that  the  sea-urchins,  or 

1  Quart.  Joitrn.  Geol.  Soc.  vol.  LI.  pp.  255 — 312  (1895). 

2  Ibid.,  pp.  306,  307. 


24  INTRODUCTORY.  [CHAP. 

echinoderms,  yield  still  more  conclusive  evidence.  "As  I  have 
previously  pointed  out,"  he  writes,  "the  intimate  affinity  between 
those  of  the  West  Indies  and  the  Mediterranean  can  only  be 
explained  by  the  assumption  of  the  existence  of  a  shallow-water 
connection  across  the  Central  Atlantic  in — at  latest — Miocene 
times.  That  the  fauna  did  not  follow  along  the  shores  of  the  North 
Atlantic  basin,  is  shown  by  its  absence  from  the  northern  Miocenes 
of  Europe  and  North  America.  The  evidence  now  adduced  from 
the  fossil  corals  of  Barbados  lends  support  to  this  view,  as  showing 
that  the  West  Indian  fauna  is  only  a  fragment  of  that  of  the 
Mediterranean  Miocene,  and  has  received  nothing  from  the 
Pacific.  This  is  in  full  agreement  with  Prof.  Suess's  theory  that 
the  Atlantic  is  of  comparatively  recent  geological  age,  and  arose 
by  the  gradual  enlargement  of  two  bays  which  ran  north  and  south 
from  a  sea  that  once  extended  across  the  Mid-Atlantic  from 
Europe  to  America,  including  both  the  Mediterranean  and  the 
Caribbean  Sea." 

The  question  of  the  southward  extension  of  America,  Africa, 
and  Australia  to  join  the  Antarctic  continent  during  Tertiary  times 
is  alluded  to  in  the  sequel. 

Summing  up  the  evidence  in  regard  to  the  permanency  of 
oceans  and  continents,  Dr  Blanford1  several  years  ago  observed 
"  that  whilst  the  general  permanence  of  ocean-basins  and  conti- 
nental areas  cannot  be  said  to  be  established  on  anything  like 
firm  proof,  the  general  evidence  in  favour  of  this  view  is  very 
strong.  But  there  is  no  evidence  whatever  in  favour  of  the 
extreme  view  accepted  by  some  physicists  and  geologists  that 
every  ocean-bed  now  more  than  1000  fathoms  deep  has  always 
been  ocean,  and  that  no  part  of  the  continental  area  has  ever  been 
beneath  the  deep  sea.  Not  only  is  there  clear  proof  that  some 
land-areas  lying  within  continental  limits  have  at  a  comparatively 
recent  date  been  submerged  over  1000  fathoms,  whilst  sea-bottoms 
now  over  1000  fathoms  deep  must  have  been  land  in  part  of  the 
Tertiary  era,  but  there  are  a  mass  of  facts  both  geological  and 
biological  in  favour  of  land-connection  having  formerly  existed  in 
certain  cases  across  what  are  now  broad  and  deep  oceans." 

1  Appendix,  No.  8,  p.  107. 


I.]  ZOOLOGICAL   REALMS.  25 

Although  much  previous  work  had  been  done  on  the  subject, 
the  first  real  attempt  to  divide  the  land-areas  of  the 

,    ,       .  i-i  •  •  Zoological 

globe  into  zoological  provinces,  or  regions,  was  made     Realms  and 
by  Dr  P.  L.  Sclater1  in  1858.     According  to  this     Regions' 
scheme,  which  was  mainly  based  on  the  study  of  Passerine  birds, 
the  world  was   parcelled   out   into   the   following  six   zoological 
regions,  viz.: — 

1.  Palczarctic ;  Europe,  Northern  Africa,  Northern  and  Cen- 
tral Asia. 

2.  Ethiopian ;  Africa  south  of  the  Atlas,  and  Madagascar. 

3.  Indian,  renamed  Oriental  by  Dr  Wallace;  India,  South- 
eastern Asia,  and  part  of  the  Malay  Archipelago. 

4.  Australian  ;  Australia,  with  New  Guinea  and  the  adjacent 
islands,  New  Zealand,  and  Polynesia. 

5.  Nearctic  ;  America  as  far  south  as  Mexico. 

6.  Neotropical ;  Central  and  South  America,  with  the  West 
Indies. 

This  scheme,  which  has  been  adopted  and  developed  in  the 
brilliant  writings  of  Dr  Wallace,  has  the  important  merit  that  it 
coincides  to  a  great  extent  with  the  leading  geographical  divisions 
of  the  globe.  It  has,  however,  the  serious  drawback  that  it  gives 
no  greater  rank  to  Australasia  and  South  America  than  to  the  other 
divisions ;  whilst  the  remarkable  difference  between  the  fauna  of 
Africa  and  Madagascar  is  overlooked.  Further,  the  northern  parts 
of  America  are  widely  separated  from  those  of  Europe  and  Asia 
to  which  they  are  faunistically  extremely  close. 

It  should  be  added  that  in  Dr  Sclater's  scheme  the  first  four 
regions,  or  those  belonging  to  the  Old  World,  were  brigaded 
together  under  the  title  of  PAL^OG^EA,  while  the  two  last,  or  New 
World  regions,  were  bracketed  as  NEOG^EA. 

The  next  important  classification  was  one  propounded  in  1868 
by  Professor  Huxley2,  who,  basing  his  conclusions  on  the  distri- 
bution of  the  game-birds,  divided  the  world  into  a  northern  and  a 
southern  division,  taking  the  name  of  ARCTOG^EA  for  the  former, 
and  NOTOG^EA  for  the  latter ;  Notogaea  being  further  sub-divided 
into  a  Novo-Zelanian  (New  Zealand),  Australian,  and  Austro- 

1  Appendix,  No.  26.  '2  Ibid.,  No.  18. 


26  INTRODUCTORY.  [CHAP. 


Columbian  region,  the  latter  being  equivalent  to  the  Neotropical 
of  Sclater. 

Six  years  later,  Dr  Sclater1,  who  had  by  this  time  turned  his 
attention  to  the  distribution  of  mammals,  proposed  to  group  the 
regions  he  had  previously  named  under  three  larger  divisions, 
making  a  fourth  division  for  New  Zealand  and  Polynesia.  This 
scheme  is  as  follows,  viz. : — 

I.  ARCTOG^EA. — Palsearctic,  Nearctic,  Oriental,  and  Ethiopian 
regions. 

II.  DENDROG^A. — Neotropical  region. 

III.  ANTARCTOG^EA. — Australian    region   (exclusive   of  New 
Zealand  and  Polynesia). 

IV.  ORNITHOG^EA. — New  Zealand  and  Polynesian  region. 

So  far  as  mammals  are  concerned,  this  scheme  was  a  great 
advance  on  the  first  one,  although  the  distinctness  of  Madagascar 
was  not  recognised,  while  the  Palasarctic  and  Nearctic  regions 
were  still  maintained.  Most  of  the  names  for  the  major  divisions 
are,  however,  open  to  objection. 

In  1878  Dr  Heilprin2,  who  does  not  employ  these  larger 
groups,  proposed,  after  a  suggestion  of  Professor  A.  Newton,  to 
unite  Dr  Sclater's  Palsearctic  and  Nearctic  regions  under  the 
common  title  of  the  Holarctic  region ;  separating,  however,  from 
the  former  a  "transitional"  Mediterranean  region,  and  from  the 
latter  a  similar  Sonoran  region. 

A  further  step  was  made  in  1890  by  Dr  Blanford8,  who  pro- 
posed the  following  scheme,  viz. : — 

I.  Australian  region. 

II.  South  American  region. 

III.  Arctogaan  region ;    this  being  divided   into    Malagasy, 
Ethiopian,  Oriental,  Aquilonian  ( —  Palaearctic  and  northern  part 
of  Nearctic),  and  Medio-Columbian  ( =  Sonoran). 

Several  other  minor  modifications  have  been  suggested  from 
mammalian  evidence,  Dr  Allen4  in  1892  reviving  the  view  that  the 
Oriental  and  Ethiopian  regions  should  be  united,  under  the  name 
of  the  Indo-African ;  but  the  next  most  important  memoir  is  that 

1  Appendix,  No.  27.  2  Ibid.,  No.  17. 

3  Ibid.,  No.  8,  p.  76.  4  Ibid.,  No.  2. 


I.]  REALMS   AND   REGIONS.  2J 

of  Dr  Hart  Merriam1  in  1892,  whose  views  are  fully  discussed  in 
the  sequel. 

It  will  accordingly  suffice  for  our  present  purpose  to  say  that  in 
1893  an  anonymous  writer2  proposed  to  take  the  terms  NOTOG^EA, 
NEOG^A,  and  ARCTOG^A  to  indicate  the  three  major  divisions  of 
Dr  Blanford's  classification ;  the  same  terms  being  used  by  Mr 
W.  L.  Sclater3  in  a  nearly  similar  sense. 

The  following  scheme  is  the  one  adopted  in  the  present 
volume,  viz. : — 

I.  NOTOG^EIC  REALM. — i.   Australian  Region. 

2.  Polynesian  Region. 

3.  Hawaiian  Region. 

4.  Austro-Malayan  Region. 

II.  NEOG^EIC  REALM. — Neotropical  Region. 

III.  ARCTOG^EIC  REALM. —  i.    Malagasy  Region. 

2.  Ethiopian  Region. 

3.  Oriental  Region. 

4.  Holarctic  Region. 

5.  Sonoran  Region. 

It  will  be  noticed  that  the  three  realms  correspond  to  the  three 
great  evolutionary  centres  of  mammals  alluded  to  in  an  earlier 
page. 

It  may  be  added  that,  in  a  work  expressly  devoted  to  the 
geographical  distribution  of  mammals,  it  will  be  unnecessary  to 
allude  to  the  schemes  proposed  on  the  evidence  of  other  groups  of 
animals,  and  we  may  accordingly  proceed  forthwith  to  the  con- 
sideration of  the  distinctive  features  of  the  various  realms  and 
regions  here  adopted. 

1  Appendix,  No.  19.         2  Ibid.,  No.  4,  p.  289.          3  Ibid.,  No.  28. 


CHAPTER   II. 

THE    NOTOG^IC   REALM. 

Definition  and  Characters  of  the  Realm — Australian  Region — Monotremes— 
Marsupials — Rodents — Carnivores — Ungulates — Bats — List  of  Australian 
and  Papuan  Genera — Polynesian  Region  —  Hawaiian  Region — Austro- 
Malayan  Region — Palaeontological  History  of  Marsupials — How  Australia 
received  its  Fauna. 

THE  term  Notogaea  was  first  proposed,  as  stated  in  the  preced- 
ing chapter,  by  Professor  Huxley1,  to  include  not  only  the 
Australian  region  of  Dr  Sclater,  but  likewise  the  Neotropical  region 
(Austro-Columbia) ;  but  an  anonymous  writer2  appears  to  have 
been  the  first  to  restrict  it  to  the  former  of  these  areas3.  This 
view,  as  being,  on  the  whole,  the  most  convenient,  is  adopted  here; 
and  the  Notogasic  realm  may  accordingly  be  taken  as  the  first  of 
the  three  primary  zoological  divisions  of  the  globe,  and  as  equiva- 
lent to  the  Australian  region  of  Drs  Sclater  and  Wallace.  Accord- 
ing to  the  latter  writer4,  "  its  central  and  most  important  masses 
consist  of  Australia  and  New  Guinea,  in  which  the  main  features 
of  the  region  are  fully  developed.  To  the  north-west  it  extends  to 
Celebes,  in  which  a  large  proportion  of  the  Australian  characters 
have  disappeared,  while  Oriental  types  are  mingled  with  them  to 
such  an  extent  that  it  is  rather  difficult  to  determine  where  to 
locate  it.  To  the  south-east  it  includes  New  Zealand,  which  is  in 
some  respects  so  peculiar  that  it  has  even  been  proposed  to  con- 
stitute it  a  distinct  region.  On  the  east  it  embraces  the  whole  of 
Oceania  [Polynesia]  to  the  Marquesas  and  Sandwich  Islands, 

1  Appendix,  No.  18.  '2  Ibid.,  No.  4. 

3  The  term  Antarctogsea  has   been  proposed   by  Dr  Sclater  (Appendix, 
No.  27,  p.  214),  for  this  area,  but  it  is  not  a  happy  one. 

4  Appendix,  No.  32,  vol.  i.,  p.  387. 


CHAP.  II.]    THE    NOTOG/EIC   REALM  :    ITS   CHARACTER.     29 

where  a  very  scanty  and  often  peculiar  fauna  must  be  affiliated  to 
the  general  Australian  type."  To  the  north-east  the  line  of  de- 
marcation of  the  realm  from  the  Oriental  region  of  Arctogsea  has 
been  finally  fixed  at  the  deep  channel  separating  the  islands  of 
Celebes  and  Lombok  on  the  one  side  from  those  of  Borneo  and 
Bali  (at  the  extremity  of  Java)  on  the  other ;  this  division  being 
now  well  known  under  the  name  of  "  Wallace's  line." 

All  writers  are,  however,  by  no  means  agreed  as  to  the  right  of 
the  whole  of  the  area  thus  indicated  to  form  a  single  zoological 
division.  Before  the  publication  of  Dr  Wallace's  great  work, 
Professor  Huxley  had  proposed  to  separate  New  Zealand  as  a 
region  of  equal  rank  with  his  Australasian  region.  At  a  later  date 
Professor  Heilprin1  suggested  that  the  Australian  realm  should 
include  only  Australia,  Tasmania,  New  Guinea,  with  the  smaller 
Papuan  islands,  and  New  Zealand ;  Polynesia,  including  all  the 
islands  lying  to  the  east  of  the  Coral  Sea,  being  raised  to  the  rank 
of  a  distinct  realm  (the  Polynesian),  while  the  Austro-Malayan 
islands  were  regarded  as  forming  a  transitional  tract  between  the 
Australian  realm  and  what  is  here  termed  the  Oriental  region.  In 
this  connection  it  may  be  well  to  notice  that  the  Austro-Malayan 
sub-region  of  Dr  Wallace  is  by  no  means  coterminous  with  the 
Austro-Malayan  transition-tract  of  Heilprin,  the  former  including, 
and  the  latter  excluding  New  Guinea. 

So  far  as  mammals  alone  are  concerned,  Notogaea  is  widely 
separated  from  the  whole  of  the  rest  of  the  world  by  being  the  sole 
habitat  (both  now  and  in  the  past)  of  the  typical  diprotodont 
marsupials  and  the  monotremes;  although  it  must  not  be  sup- 
posed that  either  of  these  groups  is  distributed  over  the  entire 
area.  As  a  matter  of  fact,  apart  from  introduced  rodents, 
Polynesia  is  devoid  of  mammalian  life  with  the  exception  of  bats 
and  a  rat 2,  while  New  Zealand  has  but  two  representatives  of  the 
former  group,  and  a  rat  which  may  or  may  not  be  indigenous. 
But  wherever  we  meet  with  a  fully  developed  mammalian  fauna, 
as  in  the  transitional  Austro-Malayan  islands,  there  a  certain 
number  of  marsupials  are  met  with,  although  the  monotremes 

1  Appendix,  No.  17. 

2  Mus  exulans,  see  Proc.  Zool.  Soc.  1895,  p.  338. 


30  THE    NOTOGyEIC    REALM.  [CHAP. 

are  restricted  to  Australia,  with  Tasmania ;  and  New  Guinea,  with 
the  adjacent  islands,  such  as  the  Aru  group. 

The  Notogaeic  realm,  as  denned  above,  may  be  conveniently 
divided  into  four  distinct  regions,  as  follows.  Firstly  the  Australian 
region,  comprising  Australia,  Tasmania,  New  Guinea  and  the  adja- 
cent Papuan  islands;  characterised  by  marsupials  and  monotremes 
forming  by  far  the  predominant  element  in  the  mammalian  fauna. 
Secondly  the  Austro-Malayan  region,  embracing  Lombok,  Celebes 
and  the  other  islands  lying  between  them  and  the  Australian  region; 
this  area  being  characterised  by  the  absence  of  monotremes  and  by 
the  marsupials  (all  of  which  belong  to  the  diprotodont  division  of 
the  order)  forming  only  a  small  minority  of  the  mammalian  fauna. 
Thirdly,  there  is  the  Hawaiian  region,  including  only  the  Sandwich 
Islands;  and,  lastly,  the  Polynesian  region,  which  maybe  taken  to 
include  all  the  islands,  save  those  last  named,  lying  to  the  eastward 
of  the  Coral  Sea,  together  with  New  Zealand,  and  is  characterised 
by  the  general  absence  of  terrestrial  mammals.  There  is  some 
difficulty  in  deciding  whether  the  islands  of  the  Solomon  group 
should  be  included  in  this  region,  or  classed  with  the  Papuan 
division  of  the  Australian  region,  seeing  that,  in  addition  to  a 
considerable  number  of  bats,  they  have  four  species  of  mice,  and 
one  diprotodont  marsupial  (Phalanger)1.  When,  however,  the 
poverty  of  this  fauna  as  compared  with  that  of  Papua  is  taken 
into  consideration,  and  it  is  also  borne  in  mind  that  Mr  C.  Hedley 2 
has  come  to  the  conclusion  that  the  Solomon  Islands,  together 
with  the  New  Hebrides,  New  Caledonia,  Fiji,  and  Norfolk  Island, 
are  closely  connected  by  means  of  their  flora  with  New  Zealand, 
and  have  but  little  in  common  with  Australia  and  New  Guinea,  it 
seems  preferable  to  include  the  former  group  in  the  Polynesian 
region.  On  the  same  grounds,  New  Zealand  is  regarded,  in 
accordance  with  the  views  of  Heilprin,  as  also  forming  a  portion 
of  the  same  zoological  region,  and  not  as  the  representative  of  a 
separate  region  by  itself.  The  fauna  of  the  Solomon  Islands  has 
doubtless  been  derived  directly  from  that  of  the  Duke  of  York 
group,  which  clearly  belongs  to  the  same  region  as  New  Guinea, 
and  shows  a  much  more  strongly  marked  Papuan  facies,  having 
three  species  of  mice,  and  four  marsupials. 

1  See  Thomas,  Appendix,  No.  30.  2  Appendix,  No.  16. 


II.]  AUSTRALIAN    REGION.  31 

Although  the  northern  half  of  Australia  lies  within  the  tropics, 
yet  few  portions  of  this  great  island  present  that 
luxuriance  of  vegetation  which  we  are  accustomed 
to  associate  with  tropical  scenery ;  and  large  tracts 
of  the  interior,  owing  doubtless  to  the  absence  of  elevated  moun- 
tain ranges  in  the  central  districts,  form  arid  sandy  deserts  more  or 
less  unsuited  to  the  maintenance  of  animal  life.  The  coast  regions 
and  the  borders  of  the  larger  rivers  are  accordingly  those  where 
vegetation  flourishes  best ;  the  finest  tracts  of  pasture-country,  well 
supplied  with  water,  lying  to  the  east  and  south-east,  and  Victoria 
possessing  a  mountain  range  whose  summits  are  perpetually 
clothed  with  snow.  Mountains  also  occur  in  the  dry  and  hot 
western  districts.  Although  Tasmania  enjoys  moister  conditions, 
Australia  as  a  whole  is  characterised  by  the  lack  of  water  and  the 
general  dryness  of  its  climate ;  and  it  is  probable  that  to  this 
aridity  the  number  of  jumping  animals,  such  as  kangaroos,  rat- 
kangaroos,  and  jerboa-rats, — now  characteristic  of  this  part  of  the 
region — is  due,  since  such  creatures  are  admirably  adapted  for 
traversing  long  distances  in  search  of  food  and  water.  On  the 
other  hand,  New  Guinea,  together  with  the  Papuan  islands,  has  a 
moist  tropical  climate,  essentially  different  from  that  of  Australia, 
but  similar  to  the  conditions  obtaining  in  a  large  portion  of 
the  Austro- Malayan  islands.  Hence  it  is  not  to  be  wondered 
at  that  the  mammals  of  New  Guinea  differ  very  markedly 
from  those  of  Australia ;  this  being  especially  noticeable  in  the 
paucity  of  typical  jumping  kangaroos,  and  the  proportionately 
large  number  of  arboreal  members  of  this  group.  Nevertheless  the 
mammalian  fauna  of  Queensland  and  North  Australia  exhibits  a 
marked  approximation  to  that  of  New  Guinea,  one  species  of 
kangaroo,  as  well  as  a  cuscus,  a  striped  phalanger  (DaJylopsila),  a 
flying  phalanger  (Petaurus),  a  pouched-mouse  (Phascologale),  and 
an  echidna,  being  common  to  the  two  areas,  and  it  is  in  these 
countries  alone  that  tree-kangaroos  are  met  with.  From  these 
resemblances  in  their  faunas — and  especially  from  the  restriction 
of  the  monotremes  to  these  two  areas, — there  can  be  no  question 
as  to  the  propriety  of  including  Australia  and  New  Guinea  in  the 
same  zoological  region,  and  thus  separating  the  latter  country  from 
the  Austro-Malayan  region. 


THE    NOTOG^IC   REALM. 


[CHAP. 


The  egg-laying  mammals,  or  monotremes,  constitute  not  only 
a    distinct    order    (Monotremata),    but    likewise    a 

Monotremes.  .    N 

separate  sub-class  (Prototheria);  and  are  broadly 
distinguished  from  all  other  members  of  their  class  by  laying  eggs, 
from  which  the  young  are  in  due  course  hatched;  as  they  are  likewise 
by  the  milk-glands  of  the  female  opening  on  the  surface  of  the 
skin  by  means  of  a  number  of  minute  perforations,  without  being 
furnished  with  nipples.  The  group  is  represented  by  three  genera, 


FIG.   i.     THE  DUCKBILL.     (Ornithorhynchus  anatinus.} 

one  of  which  is  widely  different  from  the  other  two  and  forms  a 
family  by  itself,  while  the  latter  constitute  a  second  family.  The 
duckbill  {Ornithorhynchus  anatinus},  as  the  single  representative 
of  the  first  family  (OrnithorhynchidcB)  is  commonly  termed,  is  an 
aquatic,  somewhat  mole-like,  burrowing  animal,  easily  recognised 
by  the  expansion  of  the  muzzle  into  a  broad  duck-like  beak 
covered  during  life  with  a  sensitive  skin,  and  also  by  the  broadly 
webbed  feet,  of  which  the  soles  are  naked  and  devoid  of  pads. 


II.]  MONOTREMES.  33 

Although  in  the  adult  the  mouth  is  furnished  only  with  horny 
plates,  in  young  individuals  the  sides  of  the  jaws  are  provided 
with  three  pairs  of  molar  teeth,  quite  unlike  those  of  any  other 
living  mammals.  At  the  present  day  the  duckbill  is  confined  to 
Queensland  south  of  latitude  18°,  New  South  Wales,  Victoria, 
South  Australia,  and  Tasmania;  it  is  represented  by  an  extinct 
species  from  the  Plistocene  of  Queensland,  but  otherwise  the 
pakeontological  record  of  the  group  is  a  complete  blank. 

Just  the  same  is  the  case  with  the  echidnas,  or  spiny  anteaters 
(Echidnidce],  of  which  the  only  fossil  remains  known  have  been 
obtained  from  the  superficial  deposits  of  New  South  Wales. 
Terrestrial  and  fossorial  in  their  habits,  the  echidnas  differ  from 
the  duckbill  in  having  the  muzzle  in  the  form  of  an  exceedingly 
slender  cylindrical  toothless  beak,  furnished  with  an  extensile 
worm-like  tongue ;  while  the  fur  is  thickly  mingled  with  short 
spines,  the  tail  being  rudimental,  and  the  unwebbed  toes  provided 
with  extremely  powerful  claws.  Of  the  two  species,  the  common 
five-clawed  echidna  (Echidna  aculeata]  extends  from  south-eastern 
New  Guinea  throughout  Australia  to  Tasmania;  whereas  the 
three-clawed  echidna  (Proechtdna1  bruijni}  is  restricted  to  New 
Guinea. 

With  the  exception  of  the  Plistocene  forms  already  alluded  to, 
no  fossil  monotremes  whatever  are  known  to  science.  It  is,  how- 
ever, not  improbable  that  certain  extinct  mammals  from  the 
Secondary  and  lower  Tertiary  rocks  of  Arctogaea,  commonly  termed 
Multituberculata,  which  will  be  more  fully  alluded  to  in  the  sequel, 
may  indicate  a  second  order  of  the  sub-class  Prototheria.  Both 
the  extinct  and  the  living  groups  are,  however,  of  a  highly  special- 
ised type,  so  that  the  one  cannot  apparently  be  regarded  as 
ancestral  to  the  other ;  but  if  the  presumed  distant  relationship 
between  the  two  be  substantiated,  it  will  indicate  that  we  are  to 
look  to  a  northern  origin  for  the  existing  monotremes. 

The  marsupials,  which  likewise  represent  both  a  separate 
order  (Marsupialia)  and  a  sub-class  (Metatheria) 

Marsupials. 

oy    themselves,    differ    from    the    monotremes    by 

producing  living  young,  and  by  the  milk-glands  of  the  female 

1  It  has  recently  been  proposed  to  substitute  the  term  Zaglossus,  which  is 
stated  to  be  earlier,  for  this  genus. 

L.  3 


34  THE    NOTOG^IC   REALM.  [CHAP. 

discharging  their  secretion  by  means  of  nipples.  From  the  higher 
mammals  (Eutheria)  they  are  distinguished  by  the  imperfectly 
developed  condition  of  the  newly-born  young,  and  the  absence 
of  any  prenatal  connection  between  the  vascular  system  of  the 
foetus  and  the  maternal  parent  by  means  of  the  organ  known 
as  the  placenta.  Very  generally  the  young  are  carried  about  for 
some  time  after  birth  in  a  pouch  situated  on  the  abdomen  of  the 
parent,  where  they  at  first  remain  immovably  fixed  to  the  nipples, 
the  milk  being  injected  into  their  throats  by  the  action  of  a  special 
muscle.  In  the  carnivorous  and  insectivorous  forms  the  number 
of  incisor  teeth  in  the  upper  jaw  usually  exceeds  the  three  pairs 
which  form  the  general  maximum  limit  in  the  higher  mammals.  A 
further  peculiarity  of  the  order  is  to  be  found  in  the  replacement 
of  the  teeth.  Instead  of  the  whole  or  nearly  the  whole  of  the  first, 
or  milk-set  of  teeth  in  advance  of  the  true  molars  or  hinder  cheek- 
teeth being  replaced  by  a  second  set  of  permanent  teeth,  only  one 
tooth  is  thus  (and  that  by  no  means  invariably)  replaced.  The 
tooth  thus  replaced  was  long  regarded  as  corresponding  to  the  last 
or  fourth  milk-molar  of  the  higher  mammals,  while  the  apparently 
replacing  tooth  was  identified  with  the  last  or  fourth  premolar  of 
the  same.  From  recent  researches,  however,  it  would  seem  that 
in  reality  this  is  not  a  case  of  true  replacement,  and  that  the  tooth 
which  makes  its  appearance  late  in  life  is  a  retarded  premolar, 
representing  the  fourth  in  that  series,  while  the  replacing  tooth  is 
the  fifth. 

Marsupials  may  be  divided  into  two  main  sections  or  sub- 
orders, readily  distinguished  from  one  another  by  their  dentition, 
both  of  which  are  represented  in  Notogaea.  In  the  first  of  these, 
or  Diprotodont  sub-order,  which  is  the  more  specialised  of  the  two, 
the  incisor  teeth  are  separated  by  a  gap  from  those  of  the  cheek- 
series,  and  do  not  usually  exceed  three  in  number  on  each  side  of 
the  upper  jaw1,  and  in  the  lower  jaw  are  generally  reduced  to  a  single 
pair,  while  the  tusks,  or  canines  (<:),  are  either  small  or  wanting.  In 
their  habits  the  members  of  this  section  are  more  or  less  exclusively 
herbivorous.  On  the  other  hand,  in  the  Polyprotodont  mar- 
supials, all  of  which  are  mainly  carnivorous  or  insectivorous  in 

1  The  only  exception  to  this  occurs  in  the  South-American  forms. 


II.]  MARSUPIALS.  35 

their  diet,  the  incisor  teeth  are  numerous  and  pointed,  the  canines 
are  large  and  well  developed,  and  the  whole  of  the  anterior  teeth 
form  a  series  more  or  less  nearly  continuous  with  those  on  the 
sides  of  the  jaws.  The  typical  diprotodonts,  or  those  in  which 
two  of  the  toes  of  the  hind  foot  are  enclosed  in  a  common  integu- 
ment1, are  exclusively  confined  to  the  Notogaeic  realm,  where  they 
attain  their  maximum  development  in  the  Australian  region ;  but 
the  polyprotodonts  and  an  aberrant  group  of  diprotodonts  are 
still  represented  in  the  Neogaeic  realm,  while  during  the  Secondary 
and  earlier  part  of  the  Tertiary  period  the  former  were  widely 


FIG.  2.    SKULL  OF  RAT-KANGAROO. 
(To  exhibit  Diprotodont  type  of  dentition.} 

spread  over  Arctogaea.  In  the  Australian  region  marsupials  play 
the  part  of  the  eutherians  of  other  regions,  and  show  a  remark- 
able diversity  of  external  form  and  structure,  adapting  them  to  all 
modes  of  life  with  the  exception  of  the  aquatic.  And  it  is  fairly 
evident  that  within  the  limits  of  this  region  the  diprotodonts  were 
originally  evolved  from  the  more  generalised  polyprotodonts. 

Of  the  three  existing  family  groups  into  which  the  Notogaeic  di- 
protodonts are  divided  the  first  is  the  Macropodidce,  or  the  kangaroos 
and  their  allies ;  this  being  in  some  respects  the  most  specialised 
group  of  all,  and  characterised  by  certain  peculiar  features  in  the 
skull  and  dentition.  Among  these  the  typical  genus  Macropus,  in- 
cluding the  true  kangaroos,  comprises  a  total  of  twenty-three 
species,  out  of  which  twenty  are  confined  to  Australia  and 

1  The  term  syndactylous  is  applied  to  this  type  of  foot. 

3—2 


36  THE    NOTOG^IC   REALM.  [CHAP. 

Tasmania,  one  (M.  agilis)  is  common  to  Australia  and  Queens- 
land, and  two  others  (M.  bruijni  and  M.  browni}  are  confined  to 
New  Guinea  or  the  adjacent  islands.  Of  the  six  species  of  rock- 
kangaroos  (Petrogale),  none  are  found  out  of  continental  Australia, 
and  the  same  is  true  with  regard  to  the  three  representatives  of 
the  nail-tailed  wallabies  (Onychogale),  and  likewise  with  the  three 
hare-wallabies  (Lagorchestes}.  On  the  other  hand,  the  three  kinds  of 
dorca  kangaroo  (Dorcopsis)  are  exclusively  Papuan ;  while  of  the 
climbing  tree-kangaroos  (Dendrolagus),  three  are  from  Papua 
and  two  from  Queensland.  The  single  species  of  banded  wallaby 
(Lagostrophus)  is  Australian ;  as  are  also  the  whole  of  the  rat- 
kangaroos,  forming  the  genera  Potorous,  Caloprymnus,  Bettongia, 
and  sEpyprymnus,  and  likewise  the  peculiar  musk-kangaroo 
(Hypsiprymnodon),  which  serves  to  connect  the  other  members  of 
the  family  with  the  phalangers. 

Several  of  the  existing  representatives  of  the  above-mentioned 
genera  are  found  in  a  fossil  state  in  the  cavern-deposits  of  New 
South  Wales  and  the  Plistocene  formations  of  Queensland,  in 
addition  to  which  there  are  likewise  several  extinct  representatives 
of  the  genus  Macropus,  some  of  which  considerably  exceed  the 
largest  living  forms  in  point  of  size.  The  same  formations  have 
also  yielded  the  remains  of  three  extinct  genera,  namely  Palor- 
chestes,  Procoptodon,  and  Sthenurus,  all  of  which  appear  to  have 
been  allied  to  the  wallabies,  although  some  of  the  species  were 
vastly  larger  than  any  existing  kangaroo.  Another,  but  very  im- 
perfectly known  genus  Triclis,  seems  to  have  connected  the 
musk-kangaroo  so  closely  with  the  phalangers,  that  it  is  scarcely 
possible  to  draw  any  distinction  between  these  two  families. 

In  the  family  of  the  phalangers  (Phalanger  idcz],  which 
differs  from  the  more  typical  representatives  of  the  preceding 
by  the  more  generalised  characters  of  the  skull,  teeth,  and 
limbs,  there  is  an  exclusively  Australian  form  in  the  koala, 
forming  the  sole  representative  of  the  genus  of  the  same  name. 
Of  the  five  species  of  cuscuses  (Phalanger},  one  is,  however, 
common  to  northern  Australia,  New  Guinea,  and  the  Austro- 
Malayan  Islands,  while  the  other  four  are  restricted  to  the  two 
latter  areas.  The  two  species  of  true  phalanger  (Trickosurus)  are, 
on  the  other  hand,  exclusively  Australian  ;  while  the  ring-tailed 


II.]  MARSUPIALS.  37 

phalangers,  constituting  the  genus  Pseudochirus,  are  common  to 
Australia  and  New  Guinea.  Another  exclusively  Australian  type 
is  to  be  found  in  the  taguan  flying  phalanger  (Petauroides) ;  but  of 
the  non-volant  striped  phalangers  (Dactylopsila)  one  species  is 
common  to  Queensland,  the  Aru  Islands,  and  New  Guinea,  while 
the  second  is  exclusively  Papuan.  The  true  flying  phalangers  of 
the  genus  Petaurus  include  two  Australian  species,  and  a  third, 
common  to  northern  and  eastern  Australia  and  New  Guinea  and 
the  adjacent  islands.  Leadbeater's  phalanger  (Gymnobelideus), 
which  appears  to  be  closely  related  to  the  ancestral  stock 
from  which  were  evolved  the  members  of  the  last  genus,  is 
restricted  to  Victoria ;  but  the  dormouse-phalangers  of  the  genus 


FIG.  3.     SKULL  OF  EXTINCT  PHALANGER  (Thylacoleo  carnifex). 

Dromicia  have  both  Australian  and  Papuan  representatives,  while 
the  pen-tailed  phalanger  (Distachurus)  is  exclusively  from  New 
Guinea,  and  of  the  two  pigmy  flying  phalangers  (Acrobates),  one  is 
Australian  and  the  other  Papuan.  Lastly,  the  aberrant  long- 
snouted  phalanger  ( Tarsipes),  representing  a  sub-family  by  itself, 
is  confined  to  Western  Australia.  Remains  of  species  belonging 
to  some  of  the  existing  genera  have  been  disinterred  from  the 
caves  of  New  South  Wales  and  the  Plistocene  deposits  of  Queens- 
land; while  several  more  or  less  imperfectly  known  extinct 
generic  types  have  been  described.  Among  the  latter,  by  far  the 
most  remarkable  is  Thylacoleo,  which  was  a  gigantic  phalanger 
comparable  in  size  to  a  large  leopard,  and  distinguished  by  the 
great  development  of  the  last  premolar  tooth  in  each  jaw.  The 


38  THE    NOTOG^IC   REALM.  [CHAP. 

tooth  in  question  has  an  elongated  cutting-blade,  adapted  to  work 
against  its  fellow  in  the  opposite  jaw  with  a  scissor-like  action, 
somewhat  after  the  fashion  of  the  carnassial  teeth  of  a  tiger ;  but 
the  other  cheek-teeth  were  all  relatively  small,  although  the  tusks 
were  large.  The  giant  among  the  marsupials  was  the  extinct 
Diprotodon  of  the  Australian  Plistocene,  a  creature  rivalling  in  size 
the  extinct  South  American  Megalotherium,  and  allied  on  the  one 
hand  to  the  kangaroos,  and  on  the  other  to  the  phalangers.  It 
was  not,  however,  endowed  with  the  leaping  powers  of  the  former, 
and  doubtless  walked  on  the  ground  in  the  ordinary  manner,  its 
toes  having  apparently  been  covered  with  structures  intermediate 
between  hoofs  and  nails.  Nearly  related,  but  likewise  repre- 
senting a  family  by  itself,  is  the  somewhat  smaller,  but  still 
gigantic  Nototherium,  which  in  the  conformation  of  its  limb-bones 
appears  to  approximate  to  the  wombats,  and  may  consequently 
have  been,  like  those  animals,  of  fossorial  habits. 

The  last  Notogaeic  family  of  the  Diprotodont  section  is  that  of 
the  wombats  (Phascolomyidcz\  distinguished  from  all  the  preceding 
forms  by  the  presence  of  only  a  single  pair  of  incisor  teeth  in  both 
the  upper  and  lower  jaws ;  canines  being  absent,  and  the  whole 
dentition  thus  curiously  simulating  that  of  the  rodents  among 
the  higher  mammals.  All  the  three  existing  species,  which 
are  included  in  the  single  genus  Phascolomys,  are  confined  to 
Australia  and  Tasmania;  and,  except  certain  extinct  species 
belonging  to  the  same  genus,  the  only  other  member  of  the 
family  is  the  extinct  Phascolonus  (Sceparnodoti)  from  the  Australian 
Plistocene,  distinguished  by  the  peculiarly  flattened  and  chisel- 
like  form  of  the  upper  incisor  teeth.  This,  the  only  known 
species,  attained  much  larger  dimensions  than  either  of  the 
existing  wombats. 

The  Polyprotodonts  likewise  include  three  existing  families 
found  within  the  limits  of  the  Australian  region,  none  of  the  mem- 
bers of  which  stray  either  into  the  Austro- Malay  an  or  Polynesian 
regions,  although  the  separate  family  of  the  opossums  (Didelphyidcz} 
inhabits  the  New  World.  In  the  family  of  the  bandicoots  (Pera- 
melidtz},  the  two  species  of  rabbit-bandicoot  (Peragale)  are  ex- 
clusively Australian,  whereas  the  true  bandicoots  (Perameles]  have 
both  Papuan  and  Australian  representatives ;  the  third  genus 


II.] 


MARSUPIALS. 


39 


( Chceropus\  which  includes  only  the  pig-footed  bandicoot,  being 
confined  to  Australia. 

In  the  second  family,  or  Dasyurida,  the  genus  Thyladnus  is 
now  confined  to  Tasmania,  but  it  was  represented  during  the 
Plistocene  period  on  the  Australian  mainland,  where  one  species 
is  stated  to  have  been  obtained  from  beds  of  Pliocene  age.  A 
similar  distribution  also  obtains  in  the  case  of  the  genus  Sarcophifas, 
now  represented  only  by  the  well-known  Tasmanian  devil. 


FIG.  4.     BANDED  ANTEATER.     (Myrmecobius  fasciatus.) 

Although  mainly  Australian,  the  smaller  animals  known  as  dasyures 
(Dasyurus]  have,  however,  a  single  Papuan  representative ;  while 
the  pouched  mice  (Phascologale)  are  likewise  common  to  the  two 
areas,  one  of  the  species  ranging  from  New  Guinea  to  eastern  and 
southern  Australia.  On  the  other  hand,  both  the  narrow-footed 
pouched  mice  (Sminthopsis)  and  the  jumping  pouched  mouse 
(Antechinomys]  are  exclusively  Australian.  The  same  is  the  case 


40  THE    NOTOG^IC   REALM.  [CHAP. 

with  the  aberrant  banded  anteater  (Myrmecobius),  which  although 
generally  included  in  the  Dasyuridce,  should  perhaps  form  the 
type  of  a  family  by  itself;  this  animal  differing  from  all  the  fore- 
going in  the  number,  and  also  in  the  structure  of  the  cheek-teeth, 
and  thereby  making  a  marked  approximation  to  certain  Marsupials 
of  the  Jurassic  epoch  noticed  in  the  sequel. 

Before  leaving  this  family,  it  should  be  mentioned  that  certain 
extinct  Marsupials  from  the  Tertiaries  of  Patagonia,  referred  to  in 
the  next  chapter,  seem  to  be  inseparable  from  it,  while  there  are 
strong  reasons  for  regarding  one  of  them  (Prothylacinus)  as  very 
nearly  allied  to  the  existing  genus  Thylacinus. 

The  last  of  the  Australian  families  {Notary ctidce]  of  the  sub- 
order is  represented  solely  by  the  marsupial  mole  (Notoryctes), 
from  the  sandy  deserts  of  central  South  Australia ;  this  being  the 
only  member  of  the  order  which  has  taken  to  a  subterranean  mode 
of  life.  There  are  no  extinct  Australian  genera  of  the  sub-order. 

Exclusive  of  the  bats,  the  only  other  order  of  mammals  well 
represented  in  the  Australian  region  is  that  of  the 
Rodentia,  or  Gnawing  Mammals,  which  bear,  how- 
ever, a  small  proportion  to  the  marsupials,  and  all  of  which  belong 
to  the  mouse-family  (Murtdce).  And  it  is  noteworthy  that  although 
several  of  these  belong  to  generic  types  unknown  elsewhere,  the 
whole  of  them  are  animals  of  comparatively  small  size,  so  that  it 
is  possible  that  their  ancestors  may  have  been  introduced  without 
a  direct  land-connection  with  any  other  part  of  the  world.  A 
curious  feature  in  connection  with  this  group  is  that  two  of  the 
Australian  species,  namely  Hydromys  chrysogaster  and  Musfuscipes, 
are  aquatic  in  their  habits ;  whereas,  as  we  have  seen,  none  of  the 
Australian  marsupials  are  natatorial,  although  the  duckbill  is 
eminently  so.  The  Australian  water-rat  (Hydromys),  which  is 
common  to  Australia  and  New  Guinea,  belongs  to  a  sub-family 
typically  distinguished  from  all  other  Murtdce  by  the  reduction  of 
the  molar  teeth  to  two  pairs  in  each  jaw.  While  this  animal  has 
partially  webbed  toes,  and  is  strictly  aquatic  in  its  habits,  the 
allied  Xeromys  from  Queensland  is  terrestrial,  and  approximates 
to  the  more  typical  members  of  the  family,  although  to  which 
group  is  still  uncertain.  The  only  other  representatives  of  the 
sub-family  Hydromyina  are  met  with  in  the  mountains  of  Luzon, 


II.]  RODENTS.  41 

in  the  Philippine  group,  where  there  is  one  genus  allied  to 
Hydromys,  while  other  species  have  been  assigned  to  the  genus 
Xeromys.  Whereas  the  typical  Australian  representative  of  the 
latter  has  but  two  pairs  of  molar  teeth,  one  of  the  Philippine 
forms  has  three,  thus  approximating  to  more  ordinary  murines. 
The  occurrence  of  these  Australian  types  of  rats  in  the  Philippines 
is  of  the  utmost  importance  in  respect  to  Australia  having  received 
its  mammalian  fauna  from  south-eastern  Asia. 

Of  the  typical  genus  Afus,  whose  geographical  distributional 
area  includes  the  whole  of  the  eastern  hemisphere  with  the  ex- 
ception of  Madagascar  and  many  of  the  Polynesian  islands1, 
Australia  has  upwards  of  twenty-six  representatives,  while  two 
species  occur  in  the  Duke  of  York  group,  and  others  probably  on 
the  Papuan  mainland.  One  of  the  Duke  of  York  species  (M. 
prcetor)  ranges  eastwards  into  the  Solomons,  where  three  other 
kinds  are  also  found.  The  jerboa-rats  (Conilurus*)  form  a  pe- 
culiar saltatorial  group  restricted  to  the  sandy  deserts  of  the 
mainland  of  Australia,  where  they  are  represented  by  about  a 
dozen  species  ;  while  the  broad-toothed  rat  (Mastacomys)  is  con- 
fined to  Tasmania,  although  its  fossilised  remains,  like  those  of 
the  other  genus,  are  met  with  in  the  caverns  of  New  South  Wales. 
More  nearly  allied  to  the  true  rats  and  mice,  the  mosaic-tailed  rats 
(Uromys)  inhabit  Queensland  and  the  Aru  Islands,  one  of  the 
species  from  the  former  area  also  occurring  in  the  Solomons. 
Lastly,  the  prehensile-tailed  rat  (Chiruromys],  from  the  mountains 
of  south-eastern  New  Guinea,  represents  a  genus  distinguished 
from  all  other  placental  mammals  of  the  eastern  hemisphere, 
with  the  exception  of  the  British  harvest-mouse  and  the  Oriental 
binturong,  by  the  prehensile  nature  of  its  tail. 

In  connection  with  these  rodents  it  is  important  to  observe 
that  fossil  Murida  are  unknown  from  any  part  of  the  world  earlier 
than  the  Miocene  epoch,  so  that  it  is  evident  the  living  Australian 
representatives  of  the  family  are  comparatively  recent  immigrants 
into  the  region  they  inhabit. 

1  The  Pacific  rat  (Mus  exulans)  appears  to  be  widely  distributed  in  these 
islands,  see  note  on  p.  29. 

2  Commonly  known  by  the  preoccupied  name  Hapalotis. 


42  THE    NOTOG^IC   REALM.  [CHAP. 

Much  discussion  has  taken  place  with  regard  to  the  date  of 
introduction  of  the  native  dog,  or  dingo  (  Canis  dingo] 

Carnivores.          .  ,.  .    .  -jj^i^-j. 

into  Australia,  and  it  was  long  considered  that  it 
was  imported  by  human  agency.  Seeing,  however,  that  its  remains 
have  been  found  in  association  with  those  of  extinct  kangaroos 
and  Diprotodon,  there  seems  considerable  probability  of  its  being 
an  indigenous  inhabitant  of  the  country  l. 

The  only  other  non-volant  mammal  found  in  the  Australian 

region  is   a  species  of  pig  (Sus  papuensis).     This 

animal  is,    however,    so    closely    allied  to    certain 

Malayan  species  that  it  seems  quite  possible  that  its  introduction 

may  be  due  to  human  agency. 

The  Australian  region  contains  representatives  of  all  the  families 

of  Bats  with  the  exception  of  the  Neogaeic  Phyllo- 

stomatid(z\  some  of  the  genera,  such  as  the  tube- 

nosed  bats  (Uronycteris*},  among  the  Pteropodida,  being  peculiar 

to  this  and  the  Austro-  Malayan  region,  while  others  are  more  or 

less  widely  spread,  or  even  cosmopolitan.     It  will  be  unnecessary 

to  mention  the  various  genera  by  name  ;  but  the  affinity  of  the 

Notogaeic  Chiroptera  to  those  of  Eastern  Arctogaea,  as  exemplified 

by  the  abundance  of  fruit-bats  (Pteropodidce)  and  the  absence  of 

the  PhyllostomatidcR)  is  noteworthy. 

In  the  following  synoptical  list  the  higher  groups  and  genera 
(exclusive  of  Bats)  peculiar  to  the  Notogaeic  realm 
are  printed  in  italic  type  ;  the  letters  A,  P,  and  M 


Papuan  following  the  names  respectively  indicate  that  the 

Genera. 

groups  in  question  occur  in  Australia  (inclusive  of 
Tasmania),  New  Guinea  (with  the  adjacent  Papuan  islands),  or  the 
Austro-Malayan  region  ;  extinct  groups  have  an  asterisk  prefixed. 
I.     MONOTREMATA. 
Ornithorhynchidce,  A. 
OrnithorhynchuS)  A. 
EchidnidcB,  A.  P. 

Echidna,  A.  P.      (The     living    species    common    to 

both  areas.) 
Proechidna,  P. 

1  See  Ogilby,  "Catalogue  of  Australian  Mammals,"  Sydney,  1891  —  92. 

2  This  name  replaces  the  preoccupied  Harpvia. 


II.] 


LIST   OF   MAMMALS. 


43 


II.     Marsupialia. 

i.       DlPROTODONTIA,  A.  P.   M. 


Macropodidce,  A.  P. 

Mac r opus,  A.  P. 

Petrogale,  A. 

Onychogale,  A. 

Lagorchestes,  A. 

Dorcopsis,  P. 

Dendrolagus,  A.  P. 

Lagostrophus,  A. 

Potorous,  A. 

Caloprymnus,  A. 

Bettongia,  A. 

sEpyprymnus,  A. 

Hypsiprymnodon,  A. 
*Palorchestes,  A. 
* Procoptodon,  A. 
*Sthenurus,  A. 
*Triclis,  A. 

Phalangerida,  A.  P.  M. 
Koala,  A. 

Phalanger,  A.  P.  M. 
Trichosurus,  A. 
Pseudochirus,  A.  P. 
Petauroides,  A. 
Dactylopsila,  A.  P. 
Petaurus,  A.  P.  M. 
Gymnobelideus,  A. 
Dromicia,  A.  P. 
Distcechurus,  P. 
Acrobates,  A.  P. 
Tar  sip  es,  A. 
*Thylacoleo,  A. 

*Diprotodontidcz,  A. 
* Diprotodon,  A. 


(Elsewhere  represented  only 
by  an  aberrant  group  in 
South  America.) 


44  THE    NOTOG^IC   REALM.  [CHAP. 

4.  *Nototheriid(K)  A. 

* Nototherium^  A. 

5.  Phascolomyid&i  A. 

PhascolomyS)  A. 
*Phascolonus,  A. 

ii.       POLYPROTODONTIA,  A.  P. 

1.  Peramelid&i  A.  P. 

Peragale,  A. 
Perameles,  A.  P. 
Chc&ropuS)  A. 

2.  DASYURID^E,  A.  P. 

ThyladnuS)  A. 
SarcophiluS)  A. 
Dasyurus,  A.  P. 
Phascologale,  A.  P. 
Smmthopsis,  A. 
AntechinomyS)  A. 
MyrmecobiuSj  A. 

3.  Notoryctidce,  A. 

Notoryctes,  A. 

III.  Rodentia. 

i.     MuRiDjE.     Cosmopolitan. 
Hydromys,  A.  P. 
Xeromys,  A.  and  Philippines. 
Mus,  A.  P.  M. 
Conilurus,  A. 
Mastacomys,  A. 
Uromys,  A.  P. 
Chiruromys,  P. 

IV.  Carnivora. 

CANID^:,  Cosmopolitan. 
Canis,  A.  Cosmopolitan. 

V.  Ungulata. 

SUID.E.     Throughout   Eastern    Hemisphere,    except 

Australia. 

Sus,    P.     Elsewhere    throughout   greater   part   of 
Eastern  Hemisphere. 


II.]  ITS   REGIONS.  45 

VI.     Chiroptera.     All    the   families,    with    the    exception    of 

the  Neogaeic  Phyllostomatidae,  well  re- 
presented. 

The  Polynesian  region,  as  already  said,  is  characterised  by  the 
general  absence  of  non-flying  mammals,  and  there- 
fore claims  but  little  notice  here.  The  only  mar- 
supial  occurring  within  the  region  is  a  variety  of  the 
widely-spread  grey  cuscus  (Phalanger  orientalis],  which  occurs  in 
the  Solomon  Islands,  where  four  species  of  Mus  are  likewise  met 
with.  As  the  cuscus,  together  with  one  of  the  rats,  is  also  found 
in  the  Duke  of  York  group,  it  may  be  inferred  that  the  non-volant 
mammals  of  the  Solomons  have  been  derived  from  the  latter  area. 
In  addition  to  members  of  widely-spread  types,  the  Solomons 
possess  two  peculiar  genera  of  bats. 

New  Zealand  appears  to  be  inhabited  only  by  two  peculiar 
generic  types  of  bats,  each  represented  by  a  single  species,  and  a 
rat  (Mus  maorimri},  but  whether  the  latter  is  indigenous  or  intro- 
duced appears  doubtful. 

Although  a  work  devoted  to  mammals  has  little  to  do  with  an 
area  where  the  sole  member  of  the  class  is  a  bat  of 
the  genus  Atalapha,  brief  mention  must  be  made  of     R^on*"' 
the  Sandwich  Islands,  which  from  their  bird-fauna 
are  regarded  as  entitled  to  distinction  from  the  Polynesian  region. 
Of  the  birds  of  this  area,  Mr  W.  L.  Sclater  '  writes  that  the  greater 
number  not  only  of  the  species,  but  even  of  the  genera  "are 
peculiar  and  wholly  restricted  to  these  islands.     It  is,  of  course, 
among  the  smaller  land-birds  (Passeres)  that  this  individuality  is 
most  marked;  but  even  in  the  other  groups,  where  the  distribu- 
tion is  generally  wider,  the  Hawaiian  birds  are,  in  many  cases, 
local." 

Poverty,  and  an  admixture  of  Australian  and  Malayan  types, 
with  a  very  marked  preponderance  of  the  latter,  are 
the  leading  features  in  the  mammalian  fauna  of  the     lay^n  Region. 


Austro-Malayan    region.     This  area    includes    the 

islands  lying   between  Makassar  Strait  and   the  narrow  channel 

separating  Lombok  from  Bali  on  the  west  and  the  Australian  region 

1  Appendix,  No.  28. 


46  THE    NOTOG^IC   REALM.  [CHAP. 

on  the  east.  The  largest  of  these  is  Celebes,  while  those  of  the 
Moluccan  group,  such  as  Gilolo,  Buru,  Ceram,  and  Timor-Laut, 
together  with  Timor  and  Sumbawa,  are  of  smaller  size.  Unfor- 
tunately no  complete  lists  of  the  fauna  of  these  islands,  so  far  as  I 
am  aware,  have  yet  been  published. 

Commencing  with  Timor  and  the  Moluccas,  we  find  several 
of  the  latter  group  of  islands  inhabited  by  four  species  of 
cuscus  (Phalanger\  two  of  which  are  common  to  the  Australian 
region,  while  the  third  (P.  ornatus]  is  peculiar,  and  the  fourth 
(P.  celebensis],  which  in  this  group  is  found  only  in  Sanghir  Island, 
is  an  inhabitant  of  Celebes,  where  the  other  three  are  unknown. 
The  only  other  Austro-Malayan  marsupial1  is  a  variety  of  the 
Australian  lesser  flying-phalanger  (Petaurus  breviceps\  this  variety 
ranging  eastwards  from  Gilolo  to  the  New  Britain  group.  With 
the  possible  exception  of  certain  shrews,  most  of  the  few  Moluccan 
species  of  eutherian  mammals  appear  to  be  identical  with  those 
of  Celebes,  whence  they  were  probably  introduced.  A  deer  from 
Timor  has  received  a  distinct  name  (Cervus  timoriensis\  and  the 
same  island  is  also  inhabited  by  a  common  Malayan  monkey 
(Macacus  cynomolgus),  a  palm-civet  (Paradoxurus  hermaphroditus], 
and  a  true  civet  ( Viverra  tangalungci],  the  latter  being  common  to 
the  Moluccas.  The  common  Javan  porcupine  (Hystrix  javanica), 
which  is  widely  spread  in  the  Malayan  islands,  is  also  found  in 
Timor.  There  is  likewise  a  cat  in  the  same  island,  which  although 
described  under  the  name  of  Felis  megalotis  as  a  distinct  species, 
and  regarded  by  Mr  Jentink  as  such,  has  been  identified  by 
Mr  W.  L.  Sclater  in  his  "  Catalogue  of  the  Mammalia  in  the 
Indian  Museum  "  as  a  mere  variety  of  the  domestic  species.  In 
regard  to  all  the  Timorese  forms  which  are  closely  allied  to,  or 
identical  with  well-known  Malayan  species,  it  is  necessary  to  take 
into  consideration  the  well-known  partiality  of  the  Malays  for 
taming  wild  animals  and  carrying  them  about  during  their  voyages; 
and  it  is  highly  probable  that  all  or  most  of  such  animals  found 
in  Timor  have  been  thus  introduced.  From  the  small  island  of 
Flores  Mr  Jentink  has  described  a  rat  (Mus  armandvillei),  which  is 
the  largest  member  of  its  genus. 

1  The  Kei  Islands,  like  the  Aru  group,  may  be  best  affiliated  to  Papua. 


II.] 


AUSTROMALAYAN    REGION. 


47 


In  addition  to  the  above-mentioned  cuscus,  which  appears  to 
be  its  only  marsupial,  Celebes  possesses  several  peculiar  types  of 
eutherian  mammals.  Among  these  is  a  black  and  nearly  tailless 
ape  (Cynopithecus  niger)  representing  a  genus  by  itself;  while  there 
is  also  a  species  of  macaque  (Macacus  maurus)  peculiar  to  the 
southern  portion  of  the  island.  A  species  of  the  lernuroid  tarsiers 
(Tarsius  fuscomanus)  is  found  both  in  Celebes  and  the  neighbour- 
ing islands  of  Salayer  and  Sanghir,  although  represented  by  an 


FIG.  5.     FORE  PART  OF  SKULL  OF  BABIRUSA  (Bcibirusa  alfurus], 

allied  form  in  the  Philippine  group.  In  the  Carnivora  there  is  a 
Malayan  species  of  civet  ( Viverra  tangalunga),  and  also  a  peculiar 
species  of  palm-civet  (Paradoxurus  musschenbroecki).  In  the  pig- 
tribe  the  babirusa  (Babirusa  alfurus),  characterised  by  the  extra- 
ordinary development  of  its  tusks,  is  the  sole  representative  of  a 
genus  confined  to  this  island  and  Bum;  while  scarcely  less  peculiar 
is  the  small  and  somewhat  antelope-like  buffalo  known  as  the  anoa 
(Bos  depressicornis\  which  although  allied  to  the  tamarao  (B.  mindo- 


48  THE   NOTOG^IC   REALM.  [CHAP. 

rensts)1  of  the  Philippines,  has  its  nearest  relatives  in  certain 
extinct  species  from  the  Pliocene  of  Northern  India.  There  is 
also  a  true  pig  (Sus  celebensis],  nearly  allied  to  Malayan  forms ;  as 
well  as  a  deer  forming  a  variety  of  the  widely  spread  sambar 
(Cervus  unicolor}.  Among  the  rodents,  a  rat  with  an  extremely 
long  muzzle  constitutes  a  peculiar  genus  (Echinothrix)^  and  there 
are  also  other  Muridce,  as  well  as  squirrels  (Sciurid<K),  in  addition 
to  numerous  bats,  mostly  belonging  to  Oriental  types ;  certain  of 
the  squirrels,  such  as  Sciurus  prevosti,  being  common  to  the 
Malayan  countries. 

Unfortunately  there  is  absolutely  no  palseontological  evidence 
to  help  us  in  regard  to  the  past  history  of  these  islands ;  but  from 
the  living  mammalian  fauna  we  should  be  inclined  to  place  the 
whole  area  within  the  limits  of  the  Oriental  region.  On  the  other 
hand,  a  large  number  of  the  birds  both  of  the  Moluccas  and 
Celebes  are  peculiar;  and  Australian  affinities  are  displayed  by  the 
presence  of  a  bird  of  paradise  (Semioptera)  in  Gilolo  and  Batjan, 
and  of  a  cassowary  (Casuarius)  in  Ceram.  Accordingly,  it  may  be 
well  to  include  not  only  the  Moluccas,  but  likewise  Celebes  within 
the  limits  of  the  Notogseic  realm,  which  will  then  embrace  the 
whole  of  the  countries  where  monotremes,  typical  diprotodont 
marsupials,  birds-of-paradise,  and  cassowaries  occur.  Still  it 
must  be  confessed  that  this  is,  after  all,  mainly  a  matter  of  con- 
venience, seeing  that  since,  as  will  be  shown  below,  the  diprotodont 
marsupials  have  in  all  probability  originated  in  the  Australian 
region,  those  inhabiting  the  Austro-Malayan  region  must  ap- 
parently be  regarded  as  comparatively  late  immigrants  from  the 
south-east2;  the  same  being  also  true  with  regard  to  the  single 
bird-of-paradise  and  the  cassowary  inhabiting  this  area.  And  it 
is  noteworthy  that  both  genera  of  Austro-Malayan  marsupials  are 
precisely  such  as,  from  their  arboreal  habits,  would  be  likely  to  be 
transported  on  floating  timber.  Dr  Wallace  has  suggested  that 
Celebes  has  been  separated  from  the  Oriental  region  since  the 

1  It  has  been  suggested  that  this  animal  is  a  hybrid  between  the  anoa  and 
the  Indian  buffalo. 

2  In  this  view  I  am  in  accord  with  Dr  Blanford,  who  (GeoL  Mag.  decade 
3,  vol.  IX.  p.  165,  1892)  writes  that  the  marsupials  of  Celebes  "are  probably 
of  later  introduction  than  the  mammals  with  Oriental  affinities." 


II.]  CELEBES.  49 

Miocene ;  this,  however,  is  obviously  too  early  a  date,  since  the 
only  known  allies  of  the  anoa  are  met  with  (in  common  with  the 
earliest  of  all  the  oxen)  in  the  Siwalik  Pliocene  of  northern 
India. 

In  regard  to  the  Moluccas,  Dr  Wallace1  observes  that  the 
absence  of  many  characteristic  groups  of  Papuan  birds,  and 
likewise  of  kangaroos  and  the  smaller  marsupials  of  New  Guinea, 
leads  to  the  conclusion  that  these  islands  "  cannot  be  mere  frag- 
ments of  the  old  Papuan  land,  or  they  would  certainly,  in  some 
one  or  other  of  their  large  and  fertile  islands,  have  preserved  a 
more  complete  representation  of  the  parent  fauna.  Most  of  the 
Moluccan  birds  are  very  distinct  from  the  allied  species  of  New 
Guinea;  and  this  would  imply  that  the  entrance  of  the  original 
forms  took  place  at  a  remote  period.  The  two  peculiar  genera 
with  clearly  Papuan  affinities,  show  the  same  thing.  The  casso- 
wary, found  only  in  the  large  island  of  Ceram  and  distinct  from 
any  Papuan  species,  would  however  seem  to  have  required  a  land 
connection  for  its  introduction,  almost  as  much  as  any  of  the 
larger  mammalia." 

In  another  work2,  summing  up  the  general  conclusions  with 
regard  to  the  fauna  of  Celebes,  the  same  writer  observes  that  "  we 
are  fully  justified  in  classing  it  as  an  '  anomalous  island,'  since  it 
possesses  a  small  but  very  remarkable  mammalian  fauna,  without 
ever  having  been  directly  united  [during  Tertiary  times]  with  any 
continent  or  extensive  land ;  and,  both  by  what  it  has,  and  what 
it  wants,  occupies  such  an  exactly  intermediate  position  between 
the  Oriental  and  Australian  regions  that  it  will  perhaps  ever 
remain  a  mere  matter  of  opinion  with  which  it  should  properly 
be  associated.  Forming,  as  it  does,  the  western  limit  of  such 
typical  Australian  groups  as  the  marsupials  among  mammalia, 
and  the  Trichogfassida*  and  Mdiphngidct*  among  birds,  and  being 
so  strongly  deficient  in  all  the  more  characteristic  Oriental  families 
and  genera  of  both  classes,  I  have  always  placed  it  in  the  Austra- 

1  Geographical  Distribution  of  Animals,  Vol.  I.  p.  419. 

2  Island  Life,  p.  432. 

3  Equal  LoriidfB ;  includes  the  brush-tongued  lories  and  loriquets. 

4  Honey-suckers. 

L.  4 


50  THE    NOTOG.EIC   REALM.  [CHAP. 

lian  region1;  but  it  may  perhaps  with  equal  propriety  be  left 
out  of  both  till  a  further  knowledge  of  its  geology  enables  us  to 
determine  its  early  history  with  more  precision." 

Having  now  briefly  surveyed  the  leading  features  of  the  terres- 
trial mammalian  fauna  of  the  whole  Notogaeic  realm, 

Palaeonto- 

logicai  History  and  discussed  the  relationship  of  the  mammals  of 
the  Austro-Malayan  to  those  of  the  Australian  region 
(in  the  restricted  sense  of  the  term),  we  are  in  a  position  to  enter 
upon  the  consideration  of  the  probable  past  history  of  Australia 
and  New  Guinea,  so  far  as  the  same  group  of  animals  is  con- 
cerned. Before  doing  so,  it  is,  however,  essential  to  state  what 
is  known  concerning  marsupials  from  other  regions  of  the  world. 
Here  it  may  be  premised  that  in  regard  to  Australia  mammalian 
palseontological  history  is  a  total  blank  previously  to  the  Pliocene 
epoch ;  while  apparently  but  little  is  known  even  of  that  period, 
the  great  majority  of  the  fossil  mammals  of  that  country  belonging 
to  the  Plistocene  epoch  of  the  earth's  history.  As  to  the  past 
history  of  the  mammals  of  New  Guinea,  we  know  absolutely 
nothing ;  and,  as  already  mentioned,  the  same  is  the  case  with 
regard  to  those  of  the  Austro-Malayan  islands.  This,  however,  by 
no  means  exhibits  the  whole  depth  of  our  deficiency  of  informa- 
tion. Throughout  the  whole  of  eastern  Asia,  to  say  nothing  of 
Alaska  and  western  Canada,  we  have  no  information  whatever  as 
to  mammalian  life  previous  to  the  Pliocene  era;  while  even  in 
that  period  our  sole  knowledge  relates  to  a  portion  of  northern 
India  and  China.  If,  therefore,  some  of  the  modern  types  of 
marsupials  originated  in  eastern  Asia  from  the  older  forms,  the 
blank  in  the  palaeontological  history  of  the  group  relates  to  just 
the  very  countries  where  these  animals  might  naturally  be  expected 
to  occur  during  the  Tertiary  period.  While  there  is  no  record  of 
their  existence  in  Asia,  in  Europe  fossil  Tertiary  marsupials  are 
unknown  at  a  later  date  than  the  upper  Oligocene,  and  in  North 
America  than  the  middle  Oligocene,  and  the  whole  of  those 
hitherto  discovered  belong  to  the  existing  American  group  of 
opossums.  If,  however,  we  were  to  infer  from  this  that  the  whole 
order  (with  the  exception  of  that  group)  never  existed  in  conti- 

1  Equivalent  to  the  Notogaeic  realm  of  this  work. 


II.]  PALEONTOLOGY   OF   MARSUPIALS.  51 

nental  Arctogaea  after  the  Secondary  epoch,  a  very  serious  error 
might  be  committed.  And  although  it  is  improbable  that  any 
marsupials  of  an  Australian  type  ever  existed  in  Europe  or  North 
America,  there  is  no  reason  why  they  should  not  have  occurred 
in  south-eastern  Asia. 

The  extinct  dasyurids  of  the  Patagonian  Miocene  have  been 
already  mentioned,  and  these,  together  with  another  S.  American 
group,  are  more  fully  noticed  in  the  next  chapter.  With  regard 
to  the  opossums,  it  will  suffice  to  state  that  while  they  are 
unknown  in  the  aforesaid  Patagonian  deposits,  certain  species 
occur  in  the  middle  Oligocene  White  River  beds  of  the  United 
States,  and  others  in  the  lower,  middle,  and  upper  Oligocene1  beds 
of  Europe.  Although  the  number  of  their  incisor  teeth  is  unknown, 
there  is  little  doubt  that  the  European  Oligocene  opossums2  (which 
have  been  very  generally  separated  as  Pcrathtrium\  should 
be  included  in  the  existing  genus  Didelphys.  Remains  of 
these  animals  have  been  obtained  from  the  upper  Oligocene  of 
Cournon  in  France,  from  the  middle  Oligocene  beds  of  Hordwell 
in  Hampshire,  from  the  equivalent  deposits  of  Debruge  in 
Vaucluse,  and  of  Montmartre  near  Paris,  and  likewise  from 
the  Quercy  Phosphorites  in  the  south  of  France.  With  the 
exception  of  a  peculiar  South  American  group  of  diprotodonts, 
the  remaining  fossil  mammals  which  can  be  referred  to  the 
Marsupialia  are  mainly  if  not  exclusively  confined  to  the  Second- 
ary period  ;  all  being  of  small  dimensions,  and  many  of  them 
exceedingly  minute.  While  many  of  them  evidently  died  out 
without  leaving  any  existing  descendants,  one  group  seems  to 
have  been  the  ancestral  type  from  which  the  existing  Dasyurida 
have  originated.  Among  the  former,  we  have  the  family  Tricono- 
dontidce,  as  represented  by  the  genus  Triconodon  of  the  upper 
Jurassic  of  England  and  also  by  nearly  allied  forms  from  the 
corresponding  rocks  of  the  United  States.  In  this  family  the 

1  It  may  be  well  to  mention  that  the  beds  of  St  Gerand-le-Puy,  in  France, 
which  many  writers  reckon  as  lower  Miocene,  are  here  classed  as  upper  Oligo- 
cene.    See  Lydekker,  Cat.  Foss.  Mamm.  Brit.  Mus.  Pt.  ,iv.  p.  xvii. 

2  The  existing  Didelphyida  differ  from  the  Dasyurida  in  the  presence  of  four, 
in  place  of  three,  pairs  of  incisor  teeth  in  the  lower  jaw,  and  of  five  pairs  in  the 
upper  jaw  instead  of  four. 

4—2 


52  THE    NOTOG^IC    REALM.  [CHAP. 

molar  teeth  consist  of  three  simple  compressed  trenchant  cusps 
arranged  in  a  longitudinal  line  ;  the  upper  ones  biting  on  the 
outer  side  of  those  of  the  lower  jaw.  In  the  upper  jaw  the 
number  of  teeth  is  still  unknown,  but  the  lower  jaw  carries  three 
pairs  of  incisors,  four  of  premolars,  and  either  three  or  four  of  molars, 
in  addition  to  the  tusks  or  canines,  which  are  implanted  by  two 
distinct  roots.  In  this  respect  the  latter  teeth  present  an  approxi- 


FlG.    6.       INNER    SURFACE   OF    RIGHT   HALF    OF    LOWER  JAW   OF   TriCOHodon, 
OR    ALLIED    FORM. 

mation  to  those  of  the  bandicoots,  where  the  root  of  the  canine  is 
partially  divided  by  a  longitudinal  groove.  A  second  family 
(Spalacotheriidce),  likewise  represented  in  the  upper  Jurassic  rocks 
both  of  Europe  and  the  United  States,  is  distinguished  by  the 
cusps  of  the  molars  being  arranged  in  a  triangle,  with  the  apex 
pointing  inwards  in  the  upper,  and  outwards  in  the  lower  jaw  ; 
these  teeth  being  similar  in  structure  to  those  of  the  marsupial 
mole. 

Of  more  interest  is  the  large  family  of  the  Amphitheriida, 
which  may  be  taken  to  include  a  great  number  of  forms  apparently 
agreeing  with  the  opossums  in  having  four  pairs  of  lower  incisor 
teeth.  The  lower  molars  never  consist  solely  of  three  simple 
cusps  arranged  in  a  straight  line  like  those  of  the  Triconodontida, 
or  in  a  triangle  like  those  of  the  Spalacotheriidce.  Among  these 
forms  the  genus  Phascolotherium,  from  the  lower  Jurassic  Stones- 
field  Slate  of  Oxfordshire,  appears  to  have  had  only  seven  pairs  of 
cheek-teeth ;  the  lower  molars  having  three  cusps  arranged  in  a 
longitudinal  line,  of  which  the  middle  one  is  considerably  larger 


II.] 


PALEONTOLOGY   OF   MARSUPIALS. 


53 


than  the  other  two,  while  there  are  minute  additional  cusps  at  the 
two  extremities,  and  a  distinct  ledge  at  the  base  of  the  inner  side 


Nat.  size. 


FIG.  7.     IMPERFECT  RAMUS  OF  LOWER  JAW  OF  Phascolotherium. 

of  each  tooth.  In  the  allied  Amphilestes,  from  the  same  forma- 
tion, of  which  the  imperfect  lower  jaw  is  shown  in  the  annexed 
figure,  the  molars  are  of  the  same  general  type  as  in  the  preceding, 
but  much  more  numerous,  their  total  number  being  probably 
nearly  the  same  as  in  the  next  genus. 


Nat.  size. 


FIG.  8.     IMPERFECT  RAMUS  OF  LOWER  JAW  OF  Amphilestes. 

Another  type  is  represented  by  the  genera  Amphitherium  of 
the  Stonesfield  Slate  and  Amblotherium  of  the  upper  Jurassic  of 
Dorsetshire,  in  which,  in  addition  to  the  canines,  there  are  from 
six  to  eight  molar  teeth,  four  premolars,  and  four  incisors  in  each 
half  of  the  lower  jaw.  The  lower  molars  differ  from  those  of  the 
preceding  genera,  and  thereby  resemble  the  corresponding  teeth 


54  THE    NOTOG/EIC    REALM.  [CHAP. 

of  the  opossums  and  bandicoots,  in  that  they  consist  of  an  anterior 
portion  carrying  three  cusps  in  a  triangle,  and  of  a  posterior 
moiety  or  heel.  Several  more  or  less  nearly  related  genera  have 
left  their  remains  in  the  upper  Jurassic  rocks  of  the  United  States, 
among  which  Dryolestes  may  be  specially  mentioned ;  and  it 
appears  that  in  North  America  the  group  survived  till  the  succeed- 
ing Cretaceous  epoch.  The  especial  interest  attaching  to  these 
marsupials  is  that  they,  and  they  alone,  have  molar  teeth  com- 
parable in  number,  and  to  a  certain  extent  in  structure,  with  those 
of  the  living  Australian  Myrmecobius  ;  and  there  can  be  but  little 
hesitation  in  regarding  the  latter  as  the  specially  modified  descen- 
dant of  these  ancient  forms  of  mammalian  life.  It  is,  however, 
important  to  notice  that  all  the  Jurassic  types  have  four  pairs  of 
lower  incisor  teeth,  which  are  now  retained  by  the  opossums  alone, 
having  been  reduced  in  all  the  Australian  polyprotodonts  to  three. 
Although  a  very  low  type  of  mammal  (Dromatherium)  occurs 
HOWAUS-  in  t^ie  Triassic  rocks  of  North  America,  the  fore- 
traiia  received  going  include  all  the  leading  extinct  forms  that  can 
be  included  among  the  marsupials.  During  the 
Jurassic  epoch  the  group  seems  to  have  been  widely  distributed 
over  Europe  and  North  America ;  it  is  known  to  have  existed  in 
the  latter  area  during  the  Cretaceous  epoch,  and  it  probably  also 
survived  to  that  date  in  some  part  of  the  northern  half  of-  the 
Old  World.  After  that,  our  knowledge  is  a  blank  till  we  meet 
with  the  Oligocene  opossums  of  Europe  and  North  America,  and 
the  Miocene  Patagonian  marsupials ;  so  that  as  regards  the 
Eocene  epoch  there  is  absolutely  no  record  whatever  of  the  group. 
That  Australia  received  its  original  fauna  of  polyprotodont 
marsupials  from  the  northward  may  be  regarded  as  practically 
certain ;  and  the  question  as  regards  the  Notogseic  realm  accord- 
ingly narrows  itself  to  the  approximate  date  of  the  immigration. 
On  this  point  Dr  Wallace1  writes  that  "it  was  probably  far  back 
in  the  Secondary  period  that  some  portion  of  the  Australian 
region  was  in  actual  connection  with  the  northern  continent,  and 
became  stocked  with  ancestral  forms  of  marsupials ;  but  from  that 
time  till  now  there  seems  to  have  been  no  further  land-connection, 

1  Geographical  Distribution  of  Animals,  Vol.  I.  p.  465. 


II.]  ORIGIN    OF   AUSTRALIAN    FAUNA.  55 

and  the  Australian  lands  have  thenceforward  gone  on  developing 
the  marsupial  and  monotremate  types  into  the  various  living  and 
extinct  races  we  now  find  there." 

Since  this  passage  was  written,  the  case  has  been  somewhat 
materially  altered  by  the  discovery  of  the  dasyuroid  marsupials  of 
the  Patagonian  Tertiaries ;  while  recent  researches  have  tended  to 
show  that  the  alliance  between  the  Dasyuridce  and  the  Didelphyida 
is  much  more  intimate  than  was  formerly  supposed  to  be  the 
case1.  This  being  so,  it  is  a  fairly  safe  assumption  that  both 
families  are  descended  from  a  single  common  ancestral  stock 
which,  apart  from  any  question  of  a  connection  between  Australia 
and  South  America,  can  hardly  have  originated  anywhere  than  in 
the  northern  hemisphere,  seeing  that  the  Didelphyidce  are  totally 
unknown  in  Notogaea.  There  is,  however,  as  already  stated,  no 
evidence  of  the  existence  of  opossums  before  the  Oligocene ;  and 
it  is  in  the  highest  degree  improbable  that  the  two  families  were 
differentiated  as  far  back  as  the  Jurassic,  or  even  the  Cretaceous 
epoch.  Not  improbably  polyprotodont  marsupials  survived  in 
south-eastern  Asia  till  the  early  portion  of  the  Eocene  division  of 
the  Tertiary  epoch,  and  in  this  region  both  Dasyuridce  and 
Didelphyidce.  were  differentiated.  Representatives  of  the  former 
family  soon  afterwards  found  their  way  into  Australia  and  New 
Guinea,  while  the  opossums  would  appear  to  have  dispersed 
in  one  direction  into  Europe  and  in  the  other  into  North  America, 
eventually  making  their  way  from  the  latter  country  at  a  late 
epoch  in  the  Tertiary  period  into  South  America. 

Assuming  that  the  Patagonian  dasyurids  are  more  or  less 
closely  allied  to  the  Australian  forms  (and  this  certainly  appears 
to  be  the  case),  it  may  be  taken  for  granted  that  they  have  not 
originated  independently.  From  considerations  advanced  in  the 
next  chapter,  it  is  almost  impossible  to  believe  that  they  travelled 
by  way  of  North  America ;  and  if  this  be  so,  their  only  mode  of 
migration  would  be  by  means  of  a  land-bridge  between  South 
America  and  Australia  by  way  of  the  Antarctic  continent,  or 

1  In  the  British  Museum  "Catalogue  of  Marsupials  and  Monotremes," 
p.  315  (1888),  Mr  O.  Thomas  writes  that  the  family  Didelphyidce  "is,  on  the 
whole,  very  closely  allied  to  the  Dasyurida,  from  which,  were  it  not  for  its 
isolated  geographical  position,  it  would  be  very  doubtfully  separable." 


56  THE    NOTOG;EIC    REALM.  [CHAP. 

possibly  in  a  zone  nearer  the  equator1.  Assuming  such  a  connec- 
tion to  have  existed  in  Tertiary  times  (and  there  is  no  reason  why 
it  should  not  have  existed),  it  must  either  have  taken  place 
before  the  development  of  the  diprotodonts  in  Australia,  or  must 
have  been  in  such  high  latitudes,  or  so  transitory,  as  to  permit  of 
the  passage  of  only  a  few  forms.  It  is  true  that  there  is  no 
definite  evidence  that  land  mammals  ever  existed  on  the  Antarctic 
continent,  but  during  a  recent  expedition  certain  seals  were  killed 
bearing  on  their  hides  marks  which  appeared  to  have  been  inflicted 
by  the  claws  of  a  land  carnivore.  If  this  be  substantiated  by 
future  discoveries,  it  would  be  not  only  probable,  but  essential 
that  there  should  have  been  a  Tertiary  connection  between 
'Antarctica'  and  other  lands.  With  regard  to  the  probability  that 
'Antarctica'  is  of  continental  origin,  in  summarising  what  is  known 
with  regard  to  the  geology  of  'Antarctica,'  Messrs  David  and 
Smeeth  observe  that  whether  a  continent,  or  an  archipelago  the 
islands  of  which  are  united  by  thick  sheets  of  ice,  the  southern  land 
is  considered  to  have  a  superficial  area  of  4,000,000  square  miles, 
being,  therefore,  larger  than  Australia.  A  great  chain  of  volcanoes 
has  been  described,  which  in  Victorialand  rise  over  15,000  ft. 
above  the  sea.  On  the  South  American  side  of  Antarctica  may 
be  specially  noticed  the  active  volcano  of  Bridgman,  and  the 
large  and  partially-submerged  volcano  of  Deception  Island,  with 
its  crater  over  five  miles  in  diameter,  the  wall  of  which,  built  up  of 
alternating  layers  of  ice  and  volcanic  scoriae,  rises  to  1,800  ft. 
above  the  sea.  Sedimentary  rocks  of  Eocene  age,  with  fossil  trees, 
were  discovered  in  1893  at  Seymour  Island;  and  the  French  ship 
Talisman  many  years  previously  dredged  off  the  Antarctic  conti- 
nent fragments  of  rock  containing  Gyroporella,  a  fossil  plant  very 
characteristic  of  the  Triassic  rocks  of  Europe.  Near  Laurie  Island, 
in  the  South  Orkneys,  limestone  occurs.  The  rocks  collected  by 
Mr  Borchgrevink  are  of  especial  interest  as  confirming  the  theory 
that  Antarctica  is  a  continent  rather  than  an  archipelago,  for  the 
microline-granite  with  garnet  and  tourmaline,  and  the  mica-schists 
must  have  had  a  continental  origin,  such  rocks  being  almost 
unknown  in  oceanic  islands,  but  being  of  frequent  occurrence  in 
continental  areas. 

1  See  Chapter  III. 


II.]  MARSUPIALS   IN    ASIA.  57 

With  regard  to  the  presumed  survival  of  marsupials  in  south- 
eastern Asia  till  the  Tertiary  epoch,  it  may  be  mentioned  that 
although  there  is  a  total  lack  of  knowledge  of  the  early  Tertiary 
mammals  of  Asia,  yet  there  are  not  wanting  indications  of  an 
affinity  between  the  fauna  of  the  eastern  portion  of  the  latter 
continent  and  that  of  North  America  which  points  to  a  migration 
from  a  common  centre  along  the  two  sides  of  the  Pacific; — a 
migration  which  in  the  early  Tertiary  period  received  on  the 
American  side  an  abrupt  check  by  the  sea  then  dividing  North 
and  South  America.  There  is,  for  instance,  living  in  Central  Asia 
a  species  of  deer  so  closely  allied  to  the  North  American  wapiti, 
that  it  is  a  question  whether  the  two  are  really  entitled  to  specific 
distinction ;  while  the  Chinese  alligator  has  its  nearest  living  ally 
in  the  species  inhabiting  the  Mississippi.  Another  piece  of 
evidence  is  furnished  by  the  occurrence  in  the  Tertiaries  of  the 
Balkan  Peninsula  of  remains  of  the  perissodactyle  genus  Titano- 
therium,  belonging  to  a  family  only  known  elsewhere  in  North 
America.  Quite  recently  remains  of  the  North  American  masto- 
don {Mastodon  americanus]  have  been  discovered  in  eastern 
Russia1. 

The  existence  of  such  essentially  American  types  in  Eastern 
Europe  and  Central  Asia  clearly  seems  to  point  to  a  more  intimate 
connection  between  the  faunas  of  those  regions  than  exists 
between  those  of  Western  Europe  and  North  America;  and  thus 
lends  countenance  to  the  idea  that  marsupials  may  have  lingered 
on  in  Eastern  Asia  till  long  after  the  earlier  forms  had  disappeared 
from  Western  Europe.  On  this  view,  it  is  quite  probable  that  the 
opossums  of  the  Oligocene  of  Europe  may  have  been  immigrants 
into  that  area  from  the  south-east ;  this  being  confirmed  by  the 
absence  of  the  group  from  the  Ethiopian  and  Malagasy  regions. 
As  already  said,  the  existence  of  Australian  types  of  Muridce 
in  the  Philippines  affords  a  pretty  clear  proof  that  Notogaea 
received  its  fauna  from  south-eastern  Asia,  where  types  that  had 
died  out  elsewhere  at  an  earlier  epoch  appear  to  have  survived. 
Doubtless,  however,  the  Muridce  effected  their  entrance  into 
Australia  at  a  later  epoch  than  the  marsupials. 

1  See  Pavlow,  Mem.  Ac.  St  Petersbourg,  ser.  8,  vol.  i.  pt.  3  (1894). 


58  THE   NOTOG^EIC   REALM.  [CHAP. 

The  case  of  the  ratite,  or  flightless  struthious  birds  of  Notogaea, 
as  represented  by  the  extinct  moas  (Dinornithidcz)  and  the  living 
kiwis  (Apterygidce)  of  New  Zealand,  and  the  cassowaries  and 
emeus  (Casuariidcz)  of  Papua  and  Australia,  seems  to  confirm  the 
conclusions  drawn  from  the  evidence  of  the  marsupials  as  to  the 
isolation  of  the  Notogseic  realm  not  being  so  ancient  as  supposed 
by  Dr  Wallace.  It  is  to  be  presumed  that  all  will  agree  that 
more  or  less  complete  land-connections  must  have  been  necessary 
for  the  migration  of  these  birds  ;  and  if  it  can  be  shown  that  the 
group  is  a  comparatively  modern  one,  it  cannot  but  support  the 
marsupial  evidence.  Before  proceeding  to  do  so,  allusion  must, 
however,  be  made  to  the  views  of  other  writers  as  to  the  relation- 
ships of  the  different  Notogeeic  lands. 

In  Island  Life1,  Dr  Wallace  considers  that  during  the 
Cretaceous,  and  probably  also  for  a  considerable  portion  of 
Tertiary  times,  Western  Australia  was  cut  off  by  a  deep  sea  from 
the  eastern  margin  of  the  continent,  which  was  united  with  Tas- 
mania, and  possibly  also  with  New  Guinea.  The  eastern  and 
western  islands,  he  writes,  "  would  then  differ  considerably  in 
their  vegetation  and  animal  life.  The  western  and  more  ancient 
land  already  possessed,  in  its  main  features,  the  peculiar  Australian 
flora,  and  also  the  ancestral  forms  of  its  strange  marsupial  fauna, 
both  of  which  it  had  probably  received  at  some  earlier  epoch  by 
a  temporary  union  with  the  Asiatic  continent  over  what  is  now 
the  Java  Sea.  Eastern  Australia,  on  the  other  hand,  possessed 
only  the  rudiments  of  its  existing  mixed  flora  derived  from  three 

distinct  sources The  Marsupial  fauna  had  not  yet  reached  the 

eastern  land,  which  was,  however,  occupied  in  the  north  by  some 
ancestral  struthious  birds,  which  had  entered  it  by  way  of  New 
Guinea  through  some  very  ancient  continental  extension,  and  of 
which  the  emeu,  the  cassowaries,  the  extinct  Dromor?iis  of  Queens- 
land, and  the  moas  and  kiwis  of  New  Zealand,  are  the  modified 
descendants."  He  further  concludes  that  a  large  area  of  what  is 
now  the  Tasman  Sea  was  upheaved,  and  nearly,  or  quite,  con- 
nected New  Zealand  with  Australia,  whereby  the  fauna  and  flora 
then  existing  in  Eastern  Australia  were  enabled  to  colonise  New 

1  Pages  465  et  seq. 


II.]  AUSTRALIA   AND    NEW  ZEALAND.  59 

Zealand.  Finally,  this  hypothetical  bridge  sank,  isolating  such 
forms  as  had  reached  New  Zealand,  and,  soon  after,  Eastern  and 
Western  Australia  became  connected  by  land,  and  thus  assumed 
a  homogeneous  fauna.  From  this  and  other  passages,  we  are  led 
to  conclude  that  the  author  believes  that  the  Notogaeic  flightless 
birds  immigrated,  if  not  in  Cretaceous,  at  least  in  early  Tertiary 
times1,  from  more  northern  regions. 

Dr  Wallace's  conclusions  are,  however,  challenged  by  Mr  C. 
Hedley2,  who,  from  a  study  of  the  floras  of  these  regions,  supple- 
mented by  the  distribution  of  land  molluscs,  and  recent  geological 
observations,  refuses  to  admit  that  Western  Australia  ever  pos- 
sessed a  monopoly  of  characteristic  Australian  animals  or  plants. 
Although  he  considers  the  separation  of  the  western  and  eastern 
portions  of  the  continent  by  a  Cretaceous  sea  may  be  granted, 
yet  the  land  representing  Western  Australia  was  much  smaller 
than  Wallace  supposes.  "  The  shallow  inland  Cretaceous  sea  was 
studded  with  islands,  large  and  small,  which  served  the  fauna  and 
flora  as  stepping-stones  in  their  migrations  from  west  to  east  and 
from  east  to  west."  During  a  late  era  in  the  Tertiary  epoch  he 
believes  New  Guinea  to  have  been  in  connection  with  Australia ; 
and  further  urges  "  that  an  ancient  continent,  separated  on  the 
west  from  Australia  by  the  abysses  of  the  Coral  and  of  the  Tasman 
Sea,  is  represented  by  the  Solomons,  the  Fijis,  the  New  Hebrides, 
New  Caledonia,  Lord  Howe  Island,  and  New  Zealand,  with  its 
outlying  islands — In  conclusion,  I  would  contend  that  New 
Zealand  is  associated  with  the  Solomons  and  the  New  Hebrides, 
firstly,  as  a  member  of  their  volcanic  system ;  secondly,  by  com- 
munity of  fauna  and  flora;  whereas  to  Australia  it  is  related  not 
at  all  physically,  and  to  a  foreign  and  intrusive  element  bio- 
logically ;  and  that  a  theory  which  derives  the  fauna  and  flora  of 
New  Zealand  primarily  from  these  archipelagoes  and  remotely 
from  New  Guinea,  necessitates  fewer  unproved  assumptions  than 
that  which  derives  them  from  Australia." 

To  return  to  the  ratite  birds,  it  may  be  observed  in  the  first  place 

1  If  the  immigration  into  Eastern  Australia  was  Tertiary,  what  becomes  of 
the  author's  statement  that  Australia  has  been  isolated  since  the  Secondary 
epoch  ? 

a  Appendix,  No.  i6> 


60  THE   NOTOG^IC    REALM.  [CHAP. 

that  the  giant  flightless  species  such  as  Phororhachis  and  Bron- 
tornis  of  the  Patagonian  Tertiaries  have  been  recently  shown  to 
form  a  totally  distinct  group — the  Stereornithes, — and  it  is  quite 
probable  that  the  same  may  prove  to  be  the  case  with  Gastornis, 
Dasornis,  and  Diatryma  of  the  lower  Eocene  of  the  northern 
hemisphere.  Apart  from  these,  the  earliest  known  ratites  are 
Hypselornis  of  the  Pliocene  of  India — which  appears  to  be  allied 
to  the  emeus  and  cassowaries — and  the  Australian  Dromornis, 
one  species  of  which  is  likewise  of  Pliocene  age;  all  the  other 
forms  being  Plisiocene.  Moreover,  it  is  now  tolerably  certain 
that  the  true  ratites  have  originated  from  flying  birds,  and  it  is 
therefore  highly  probable  that  the  group  is  an  essentially  modern 
one1.  Accordingly,  there  is  a  strong  presumption  that  the  ances- 
tors of  these  birds  did  not  enter  Notogsea  till  comparatively  late 
in  the  Tertiary  period ;  and  that,  in  fact,  their  southern  migration 
was  not  far  removed  in  time  from  that  of  the  giant  land-tortoises, 
noticed  in  the  next  chapter.  Possibly  they  may  have  entered 
Australia  by  way  of  New  Guinea  during  the  connection  which 
Mr  Hedley  believes  to  have  existed  between  those  two  countries 
late  in  the  Tertiary  epoch ;  while  the  New  Zealand  forms  may 
have  made  their  way  by  means  of  the  presumed  land-connection 
between  these  islands  and  the  Solomons,  New  Hebrides,  etc. 

Of  course  there  is  the  difficulty  as  to  why  mammals  did  not 
enter  the  Polynesian  region  at  the  same  time;  but  it  is  conceivable 
that  even  at  this  date  the  mammalian  fauna  of  South-eastern  Asia 
may  have  been  very  poor  in  Eutherians,  while  it  is  quite  possible 
that  the  ancestral  forms  of  these  birds  may  not  have  required  the 
complete  land-connection  necessary  for  the  passage  of  the  higher 
mammals.  Such  connection  as  served  for  these  birds,  however, 
may  have  well  sufficed  for  the  transit  of  the  ancestors  of  the 
Australian  murine  rodents,  which  almost  certainly  entered  the 
country  at  a  later  date  than  the  original  marsupials  and  mono- 
tremes. 

Assuming,  then,  that  the  marsupials  and  monotremes  of  the 
Australian  region  did  not  reach  their  present  home  till  the  early 
part  of  the  Tertiary  epoch,  we  must  make  the  further  assumption 

1  Captain  Hutton  is  of  opinion  that  the  moas  originated  directly  from  flying 
birds  in  New  Zealand,  but  the  evidence  in  favour  of  this  view  appears  insufficient. 


II.]  ORIGIN    OF   MONOTREMES.  6l 

that  at  this  period  South-eastern  Asia  was  entirely,  or  to  a  great 
extent,  devoid  of  higher  mammals.  Nor  is  this  unlikely,  seeing 
that  the  ungulates  and  carnivores  of  the  lower  Eocene  of  the 
northern  hemisphere  would  clearly  have  required  time  to  spread 
themselves  to  the  southward.  Hence  it  may  be  suggested  that, 
towards  the  close  of  the  Cretaceous  epoch  there  was  first  a  migra- 
tion towards  the  south-east  of  the  ancestral  marsupials  (and 
monotremes)  inhabiting  the  northern  hemisphere  during  the 
Secondary  epoch ;  and  that  similar  migrations  of  the  higher  mam- 
mals took  place  during  Tertiary  times. 

When  once  the  ancestral  polyprotodont  marsupials  obtained  a 
footing  in  New  Guinea  and  Australia,  where  they  have  since  been 
isolated  from  any  serious  competition  with  the  higher  mammals, 
they  flourished  and  developed  to  a  degree  which  they  could  not 
possibly  have  attained  in  any  other  part  of  the  world  under 
existing  conditions.  And  it  is  doubtless  within  this  region  that 
the  more  specialised  diprotodont  types  were  evolved.  Remark- 
able as  it  undoubtedly  is,  the  present  state  of  development  of  the 
Australian  marsupials  is  nothing  to  what  it  was  during  the 
Plistocene  period,  when  there  lived  the  giant  kangaroos,  pha- 
langers,  wombats,  diprotodons,  and  nototheres  already  alluded  to, 
by  the  side  of  which  the  largest  existing  species  would  appear 
almost  dwarfs.  The  cause  of  this  universal  extinction  (for  uni- 
versal it  is)  of  all  the  larger  types  of  mammalian  life  throughout 
the  world  soon  after  the  appearance  of  man,  is  one  of  those 
problems  which  at  present  is  not  capable  of  being  satisfactorily 
solved,  as  not  even  a  glacial  period  could  have  made  a  clean 
sweep  of  the  whole  globe.  It  may  be  added  that  the  evolution  of 
the  diprotodont  marsupials  within  the  limits  of  the  Australian 
region,  points  to  the  conclusion  that  the  outlying  discuses  of  the 
Austro-Malayan  regions  are  immigrants  from  the  south-east. 

With  regard  to  the  monotremes,  it  has  already  been  mentioned 
that  there  is  no  record  of  their  past  history  beyond  the  limits  of 
the  Australian  region.  It  can,  however,  scarcely  be  doubted  that 
their  ancestors  came  from  the  north  with  the  primitive  marsupials ; 
and  if,  as  is  not  improbable,  the  Secondary  and  early  Tertiary 
Multituberculata  of  the  northern  hemisphere  are  an  allied  type, 
there  can  be  no  doubt  whatever  as  to  this  having  been  the  case. 


62  THE   NOTOG^IC   REALM.  [CHAP. 

That  Notogaea,  as  typified  by  the  Australian  region,  is  entitled 
to  form  one  of  the  three  primary  zoological  divisions  of  the  globe, 
the  distinctness  of  its  mammalian  fauna  from  that  of  any  other 
area,  not  only  at  the  present  day,  but  likewise  during  the  Plisto- 
cene,  and  probably  also  the  Pliocene  epoch,  amply  demonstrates. 
The  inclusion  within  the  same  realm  of  the  Polynesian  region, — 
which  evidently  never  had  such  a  close  connection  with  south- 
eastern Asia  during  the  time  that  area  was  mainly  populated  with 
marsupials, — is  justified  partly  on  account  of  its  containing  more 
or  less  similar  types  of  birds,  and  partly  by  the  practical  absence 
of  terrestrial  mammals.  On  the  other  hand,  the  Austro-Malayan 
region,  which  is  really  a  kind  of  zoological  No-man's-land,  is 
placed  within  the  limits  of  the  same  great  realm  more  as  a  matter 
of  convenience  than  anything  else,  although  it  is  undoubtedly 
sharply  differentiated  from  the  Oriental  region  by  Wallace's  line. 

In  conclusion,  a  few  lines  may  be  devoted  to  showing  that 
certain  other  groups  indicate  that  the  vertebrate  fauna  of  Notogaea, 
as  a  whole,  has  had  a  northern  origin.  Among  the  lizards,  the 
family  of  iguanas  (IguanidcB],  which  in  this  realm  occurs  only  in 
the  Fiji  and  Friendly  Islands,  is  represented  in  a  fossil  state  in 
the  Oligocene  beds  of  France ;  while  the  gigantic  extinct  monitor 
( Varanus  priscus)  of  the  Australian  Plistocene  appears  to  have  its 
nearest  ally  in  the  smaller  V.  sivalensis  of  the  Pliocene  of  northern 
India.  The  Notogseic  Chelonians,  which  are  confined  to  Australia 
and  New  Guinea,  all  belong  to  the  side-necked  group  (Pleurodira) 
of  the  order,  and  are  represented  by  the  families  Chelyidce.  and 
Carettochelyidce,  the  latter  containing  only  a  single  species  from 
the  Fly  River.  Now,  although  none  of  the  Australian  genera  have 
been  detected  in  the  northern  hemisphere,  the  side-necked  chelo- 
nians,  as  shown  in  the  next  chapter,  were  abundantly  represented 
there  during  the  early  Tertiary  and  Secondary  epochs ;  and  it 
is  a  remarkable  fact  that  an  extinct  genus  believed  to  be  allied 
to  Carettochelys  occurs  in  the  Eocene  of  northern  India.  Although 
from  their  aquatic,  and  sometimes  partially  marine  habits,  the 
crocodiles  are  of  less  importance  than  some  other  groups  from  a 
distributional  point  of  view,  yet  it  is  noteworthy  that  the  single 
representative  of  that  group  ( Crocodilus  porosus]  inhabiting  Noto- 
gaea (where  it  is  found  in  North  Australia,  the  Solomons,  and 


II.]  SURVIVAL   OF   OLD   FORMS.  63 

Fiji)  is  spread  over  India,  Ceylon,  and  the  south  of  China ;  while 
the  absence  of  caimans  and  jacaras  from  Notogaea  aifords,  so  far 
as  it  goes,  an  additional  argument  that  any  land  connection  which 
may  have  existed  in  Tertiary  times  between  Australia  and  South 
America  must  either  have  been  very  transitory,  or  must  have  been 
situated  in  such  latitudes  that  tropical  forms  could  not  have  used 
it  as  a  means  of  transit. 

Of  more  importance  than  all  is  the  tuatera  lizard  (Sphenodon)  of 
New  Zealand — the  sole  existing  representative  of  the  order 
Rhynchocephalia, — since  this  curious  creature  is  closely  allied  to 
the  extinct  Rhynchosanrus  and  Hyperodapedon  of  Triassic  strata  of 
the  northern  portion  of  the  Old  World.  Finally,  the  Port  Jackson 
shark  (Cestracion  philippi)  belongs  to  a  genus  which  was  living  in 
the  seas  of  Europe  during  the  Jurassic  and  Cretaceous  periods ; 
and  the  sole  living  survivor  of  the.  swarms  of  species  of  the 
Molluscan  genus  Trigonia  inhabiting  the  same  seas  occurs  in 
Australian  waters. 


CHAPTER   III. 


THE    NEOG^IC    REALM. 

Extent  and  Characters — Mammaliferous  Deposits — Monkeys — Bats — Insecti- 
vora  —  Carnivores —  Ungulates —  Horses —  Litopterna — Astrapotheria — 
Toxodonts  —  Pyrotheria — Proboscideans — Rodents — Edentates — Arma- 
dillos and  Glyptodonts — Sloths — Anteaters- — Ground-sloths — Marsupials — 
Cetaceans — Early  Distinction  of  the  Neogaeic  Fauna — Early  Separation 
of  N.  and  S.  America — Incursion  of  Northern  Mammals — Distinctness  of 
the  existing  Fauna — Origin  of  the  Santa  Cruz  Fauna — Antarctica  and  the 
South  American  element  in  the  Ethiopean  Fauna — Conclusion  —  Sub- 
regions. 

THE  second  primary  zoological  division  of  the  globe  may  be 
known  as  Neogsea1,  or  the  Neogseic  realm.  It 
characters"**  includes  only  the  Neotropical  region.  Comprising 
not  only  the  whole  of  South  and  Central  America, 
as  well  as  the  West  Indian  Islands,  this  area  also  embraces  the 
lowlands  lying  on  either  side  of  the  Mexican  plateau — the  so- 
called  tierras  calientes — thus  running  up  in  a  fork-like  manner  to 
the  lower  extremity  of  North  America.  While,  therefore,  the 
greater  part  of  this  vast  area  is  sharply  delineated  by  its  coast- 
boundary,  to  the  north  it  has  a  kind  of  No-man's-land  connecting 
it  with  the  Sonoran  region  of  Arctogaea,  and,  as  will  be  shown 
later,  through  this  transitional  area  there  has  been  a  certain 
amount  of  intermixture  of  the  proper  faunas  of  the  Neotropical 
and  Sonoran  regions.  Neogaea,  as  a  whole,  may  be  characterised 
as  a  country  of  extensive  tropical  forests  or  open  grassy  plains  ; 
deserts  occupying  only  a  few  scattered  areas  in  the  upper  Argen- 
tine (Tucuman,  etc.),  and  certain  parts  of  the  coasts  of  Chili 

1  This  term  was  originally  proposed  by  Dr  Sclater  to  include  the  whole  of 
the  New  World,  but  has  been  used  by  an  anonymous  writer  (Appendix,  No.  4) 
in  the  present  sense.  Dr  Sclater 's  term  Dendroggea  (Appendix,  No.  27, 
p.  214)  is  open  to  considerable  objection,  as  the  greater  part  of  Argentina  is 
woodless. 


CHAP.  III.]  EXTENT   AND   CHARACTERS.  6$ 

and  Peru ;  the  whole  of  the  rest  of  the  area,  with  the  exception  of 
the  higher  regions  of  the  Andes,  being  thus  admirably  adapted  for 
the  support  of  animal  life.  At  least  one  half  of  the  whole  area  is 
occupied  by  a  dense  tropical  forest,  attaining  its  richest  develop- 
ment in  the  hot  steamy  tracts  of  Brazil  and  Paraguay,  and  being 
unequalled  in  extent  in  any  other  part  of  the  globe.  With  a  width 
of  some  three  thousand  miles  from  the  Atlantic  seaboard  at  Per- 
nambuco  to  the  foot  of  the  Andes,  this  forest  extends  north  and 
south  for  nearly  thirty  degrees  of  latitude ;  while  not  only  does  it 
clothe  the  lowlands  and  valleys,  but  extends  high  up  the  mountain- 
sides, as  may  be  seen  in  the  exquisitely  lovely  harbour  of  Rio  de 
Janeiro,  where  the  forest-vegetation  commences  immediately  above 
the  wash  of  the  waves,  and  thence  extends  in  one  continuous  leafy 
mass  to  the  summits  of  mountain-ranges  at  an  elevation  of  eight 
or  nine  thousand  feet.  In  the  northern  part  of  the  area  open 
grass-lands,  like  the  "campos"  of  Brazil  and  the  savannas  of 
Venezuela,  alternate  with  the  forest ;  while  in  the  neighbourhood 
of  Buenos  Aires  the  open  pampas1  forms  one  extensive  sea  of  grass. 
The  Andes,  constituting  the  backbone  of  the  country,  run  in  one 
continuous  chain  from  north  to  south  on  the  Pacific  seaboard,  and 
present  the  usual  varieties  of  climate  and  physical  conditions 
common  to  other  elevated  mountain-ranges.  Such  climatic 
variations  are,  however,  only  an  epitome  of  those  met  with  in 
travelling  from  the  northern  to  the  southern  extremity  of  the  area ; 
the  steamy  valley  of  the  Amazons  having  a  tropical  climate,  whereas 
when  we  reach  the  southern  point  of  Patagonia  and  Tierra  del 
Fuego  we  are  in  the  midst  of  snows  and  glaciers.  To  the  hot 
forest-regions  are  restricted  the  monkeys,  marmosets,  sloths,  ant- 
eaters,  and  tree-porcupines ;  while  the  open  plains  of  the  south  are 
tenanted  by  guanaco,  deer,  viscachas,  and  rheas.  Moreover,  in 
the  forests,  the  variety  of  mammalian  life,  especially  as  regards  the 
larger  forms,  is  in  marked  contrast  to  its  comparative  paucity  in 
the  open  plains ;  not  but  that,  till  civilised  man  made  his  appear- 
ance on  the  scene,  the  number  of  individuals  may  have  been 
nearly,  if  not  quite  as  large  in  the  latter  area  as  in  an  equal  extent 
of  the  former. 

1  Although  in  Spanish  the  term  'pampas'  is  plural,  in  English  it  seems 
preferable  to  use  it  as  singular. 

L.  5 


66  THE    NEOG^IC    REALM.  [CHAP. 

At  the  present  day  the  mammalian  fauna  of  Neogaea  is  mark- 
edly distinct  not  only  from  that  of  Notogaea  but  likewise  from  that 
of  the  whole  of  the  rest  of  the  globe  (Arctogsea),  although  the  dis- 
tinction is  now,  owing  to  free  communication  with  the  north, 
much  less  marked  than  it  was  in  Tertiary  times,  and  it  is  accord- 
ingly essential  to  enter  at  once  into  the  consideration  of  the 
extinct  forms  in  order  to  show  why  this  part  of  the  world  is 
entitled  to  rank  as  one  of  three  primary  zoological  regions. 

There  are  several  districts  in  South  America  where  fossil 
Mammal-  remains  of  mammals  have  been  found ;  most  of  these 
iferous  being  remarkable  for  the  extraordinary  profusion  in 

which  the  bones  occur.  The  first  that  may  be  men- 
tioned are  the  celebrated  caves  of  Lagoa  Santa,  in  the  province  of 
Minas  Geraes,  to  the  northward  of  Rio,  which  have  yielded  re- 
mains of  a  great  variety  of  Plistocene  genera  and  species,  inclusive 
of  those  of  man.  Probably  contemporaneous  with  these  are  the 
sand-dunes  on  the  coast  of  Buenos  Aires,  which  likewise  contain 
human  remains  in  association  with  those  of  extinct  mammals ; 
while  the  so-called  Pampean  beds  of  the  Argentine  pampas  are 
apparently  somewhat  older,  although  still  pertaining  to  the  Plisto- 
cene period.  As  these  Pampean  deposits  are  exceedingly  rich  in 
fossil  mammals,  they  may  be  described  in  some  detail.  They 
form  the  great  level  tract  of  country  extending  southwards  from 
the  Rio  de  la  Plata  and  the  Parana  to  the  Rio  Colorado,  south  of 
Bahia  Blanca,  and  westwards  from  the  Atlantic  seaboard  about 
half  the  distance  to  the  Andes;  thus  occupying  some  200,000 
square  miles  of  country.  The  pampas  is  an  almost  level  grass- 
covered  plain,  intersected  by  water-courses,  and  penetrated  near 
its  margins  by  small  mountain-ranges,  while  it  is  almost  entirely 
barren  of  trees.  It  is  composed  of  a-  rich  black  alluvial  mud, 
mingled  with  beds  of  sand,  and  underlain  by,  or  in  some  places 
interstratified  with  layers  of  a  hard  white  calcareous  deposit 
known  as  tosca\  but  in  certain  spots  it  contains  beds  of  marine 
shells  belonging  to  species  still  living  in  the  adjacent  seas.  Except 
in  those  spots  where  the  tosca  comes  to  the  surface,  there  is  not  a 
stone  or  a  pebble  to  be  seen  in  the  whole  deposit,  and  near 
Buenos  Aires  the  formation  has  been  bored  to  a  depth  of  ninety 
feet  without  touching  bottom.  From  its  composition  it  is  evident 


III.]  MAMMALIFEROUS   DEPOSITS.  67 

that  the  deposit  has  been  carried  down  from  the  interior  of  the 
north  by  the  Parana,  Paraguay,  and  other  tributaries  of  what  is 
now  the  Rio  de  la  Plata ;  but  since  there  is  no  splitting  of  the 
latter  river  at  its  estuary,  it  is  evident  that  the  formation  cannot 
properly  be  called  a  delta.  That  it  is  mainly  of  freshwater  origin 
seems  evident  not  only  from  its  intrinsic  character,  but  likewise 
from  the  vast  number  of  entire  skeletons  of  mammals  buried 
within  it,  since  these  creatures  must  certainly  have  lived  very  near 
to  the  places  where  their  bones  are  now  entombed.  In  the  more 
southern  part  of  its  area  the  pampas  is,  however,  probably  to 
a  large  extent  of  estuarine  origin  ;  and  the  presence  of  layers  of 
marine  shells  in  its  uppermost  horizon  near  Buenos  Aires  proves 
that  at  least  a  portion  was  submerged  beneath  the  sea  before  its 
final  upheaval.  Whereas  the  Rio  de  la  Plata  now  flows  in  a  single 
channel  in  a  south-easterly  direction  near  the  northern  limit  of  the 
coast-portion  of  the  pampas,  it  would  seem  probable  that  the 
Parana  and  Paraguay  rivers  may  have  originally  continued  their 
southerly  course  across  the  southern  pampas,  through  which  they 
may  have  flowed  in  a  number  of  streams.  Most  likely  the  Pam- 
pean  formation  was  laid  down  in  a  slowly  subsiding  area,  in  which 
the  rate  of  deposition  approximately  counterbalanced  the  sinking, 
so  that  the  greater  part  of  it  has  been  always  land  until  the  period 
of  the  great  submergence.  After  the  latter,  the  entire  area  was 
upheaved  to  a  small  degree  above  the  sea-level,  when  the  rivers 
assumed  their  present  approximate  courses.  Whereas  in  certain 
localities  the  deposit  is  barren  of  mammalian  remains,  in  other 
spots  it  appears  absolutely  crowded  with  them,  and  the  number  of 
entire  skeletons  that  are  entombed  in  it  must  doubtless  be  counted 
by  thousands. 

Somewhat  older  than  the  Pampean  is  a  mammaliferous  deposit 
occurring  on  the  coast  near  Bahia  Blanca  in  a  small  hill  known  as 
Monte  Hermoso  ;  and  beds  of  approximately  equivalent  age  occur 
in  Catamarca  at  the  foot  of  the  Andes.  Probably  these  beds  should 
be  regarded  as  of  Pliocene  age ;  and  it  may  be  mentioned  that 
equivalents  of  these  deposits  are  met  with  in  other  parts  of 
Argentina,  while  representatives  of  the  Pampean  occur  in  Pata- 
gonia, Chili,  Bolivia,  etc.  Still  older  than  the  Monte  Hermoso 
deposits  are  the  Santa  Cruz  beds  of  Patagonia,  occurring  not  only 

5—2 


68  THE    NEOG^IC   REALM.  [CHAP. 

on  the  river  of  that  name,  but  likewise  further  north  in  the  Chubat 
district.  These  Santa  Cruz  beds  are  exceedingly  rich  in  mamma- 
lian remains,  which  are  stained  of  a  deep  black  colour ;  and  while 
they  contain  many  groups  common  to  the  Pampean  formation, 
they  lack  those  forms  found  in  the  latter  which  have  an  Arctogseic 
fades,  thus  indicating  that  we  have  reached  an  epoch  when  the 
fauna  of  South  America  was  far  more  completely  isolated  from 
that  of  the  rest  of  the  world  than  is  at  present  the  case.  By  Dr 
Ameghino  the  Santa  Cruz  beds  were  at  first  correlated  with  the 
lower  Eocene  of  Europe,  although  he  subsequently  admitted  that 
they  must  occupy  a  somewhat  higher  position  in  that  period1. 
From  the  nature  of  their  entombed  mammals2  it  is,  however, 
certain  that  they  must  be  still  newer,  and  they  cannot  be  regarded 
as  older  than  lower  Miocene.  Indeed,  they  are  underlain  by  the 
so-called  Patagonian  beds3,  which  are  of  marine  origin,  and 
contain  numerous  cetaceans,  among  which  are  whalebone  whales 
(Mystacoceti).  From  equivalent  beds  in  the  Chubat  district  of 
Patagonia,  a  large  number  of  such  cetaceans  have  been  described 
by  the  present  writer4,  and  it  is  quite  evident  that  the  oldest  age 
that  can  be  assigned  to  these  beds  is  upper  Oligocene,  seeing  that 
in  Europe  whalebone  whales  are  unknown  till  a  considerably  later 
epoch.  Probably  the  freshwater  beds  containing  the  peculiar 
mammal  alluded  to  below  under  the  name  of  Pyrotherium  are  the 
freshwater  equivalents  of  the  Patagonian  horizon5. 

Additional  evidence  in  support  of  the  comparatively  late  age 
of  the  Patagonian  beds  is  afforded  by  the  researches  of  Prof.  Cope6 
on  the  cetaceans  of  the  Miocene  (or  Upper  Oligocene?)  of  the 
United  States.  From  these  beds  have  been  obtained  remains  of 
Hypocetus  (Paracetus) — a  genus  elsewhere  known  only  from  the 
Patagonian  beds — together  with  Cetotherium  (also  occurring  in  the 
latter  deposits),  Balcenoptera,  and  a  species  of  Balcena  identified 
with  one  from  the  European  Pliocene ;  the  North  and  South 
American  species  of  Hypocetus  being  closely  allied. 

1  Bol.  Ac,  Cordoba,  Vol.  xin.  p.  260  (1894). 

2  Remains  of  the  existing  genus  Dasypus  occur  in  these  beds. 

3  Ameghino,  loc.  cit. 

4  Ann.  Mus.  La  Plata,— Pal.  Argent.  Pt.  II.  (1893). 

5  Ameghino,  op.  cit.  p.  262. 

6  Prof.  Amcr.  Phil.  Soc.  Vol.  XXXIV.  pp.  135 — 155  (1895). 


III.]  MONKEYS.  69 

With  these  preliminary  considerations,  we  pass  to  a  critical 
examination  of  the  recent  and  fossil  mammalian  fauna  of  Neogaea, 
which  will  show  how  intimately  investigations  of  this  nature  are 
connected  with  the  present  and  past  configuration  of  the  land- 
areas  of  the  globe. 

Like  as  are  many  of  them  superficially  to  their  cousins  of  the 
Old  World,  the  monkeys  of  the  New  World,  which 

,,         ,  ,  ....  .  -          Monkeys. 

are  now  confined  to  the  tropical  forest-regions  of 
the  Neogaeic  realm,  are  structurally  quite  different  to  the  former ; 
and  this  circumstance,  coupled  with  their  isolated  distribution,  in- 
dicates that  their  relationship  is,  at  most,  but  a  very  distant  one. 
Indeed  it  is  not  improbable  that  the  Old  and  New  World  monkeys 
may  have  originated  independently  from  the  group  of  lemurs,  which 
were  formerly  very  widely  distributed  over  Arctogaea.  Of  the 
two  families  of  Neotropical  monkeys,  the  first  is  represented  by 
the  beautiful  little  marmosets,  constituting  the  family  Hapalida. 
Although  having  the  same  number  (32)  of  teeth  as  the  Old  World 
monkeys,  the  marmosets  differ  in  that  the  number  of  premolars  on 
each  side  of  both  jaws  is  three,  and  that  of  the  molars  two ;  these 
numbers  being  transposed  in  the  other  group.  The  marmosets 
are  further  distinguished  by  the  broad  septum  between  the  nostrils, 
the  absence  both  of  pouches  in  the  cheeks  and  of  callosities  on 
the  buttocks,  and  the  want  of  any  prehensile  power  in  the  tail ;  in 
addition  to  which  it  may  be  noticed  that  the  thumb  is  not  op- 
posable  to  the  other  fingers,  while  all  the  digits,  with  the 
exception  of  the  first  on  the  hind  foot,  are  provided  with  long 
claws.  None  of  these  animals  are  known  from  the  Santa  Cruz 


FlG.    9.      PART    OF    RIGHT    LOWER   JAW    OF    HoniUHCulus. 

deposits.     In  the  second  family  or  Cebidce,  which  includes  much 
larger  species  than  the  diminutive  marmosets,  the  total  number  of 


70  THE    NEOG^EIC   REALM.  [CHAP. 

teeth  is  36,  owing  to  the  retention  of  the  three  pairs  of  molars 
found  in  the  Old  World  group  ;  these  monkeys  being  further  dis- 
tinguished from  the  marmosets  by  the  presence  of  nails  on  all  the 
fingers,  and  by  the  tail  being  frequently  prehensile.  The  Santa 
Cruz  beds  have  yielded  remains  of  monkeys  (JFTomunculus)  refer- 
able to  this  family,  showing  that  they  belong  to  the  original  South 
American  fauna,  but  beyond  this  nothing  is  known  as  to  the 
palseontological  history  or  origin  of  the  group.  Lemurs,  both  in 
the  past  and  the  present,  are  quite  unknown  in  the  realm  under 
consideration. 

Although  bats  might  be  thought  of  comparatively  little  im- 
portance from  a  distributional  point  of  view,  yet  the 

Bats.  :_ 

Neogseic  realm  presents  some  very  remarkable 
features  in  this  respect.  In  the  first  place  the  two  Old  World 
families  of  fruit-bats  (Pteropodtdce)  and  horse-shoe  bats  (Rhinolo- 
phidce]  are  entirely  wanting,  whereas  the  great  family  of  vampire- 
bats  (Phyllostomatida)  is  mainly  restricted  to  it,  although  a  few 
representatives  straggle  northwards  along  the  Pacific  coast  of 
North  America.  The  family  of  Emballonuridtz  is  also  more 
strongly  represented  here  than  in  any  other  part  of  the  world. 
Probably  on  account  of  their  small  size,  there  is  no  palaeonto- 
logical  record  of  the  South  American  bats  from  the  earlier 
Tertiary  deposits. 

The  Insectivora  are  almost  unrepresented  in  the  continental 
portion    of  the   realm,    although   a   shrew   (Sorex) 

Insectivora.  '  °  . 

reaches  Guatemala  and  Costa  Rica,  and  a  member 
of  the  allied  N.  American  genus  Blarina  is  also  found  in  the 
last-named  country;  both  these  being  doubtless  very  recent 
immigrants  from  the  north.  Very  remarkable  is  the  occurrence  in 
the  West  Indian  Islands  of  the  two  species  of  Solenodon,  consti- 
tuting a  distinct  family  (Solenodontidce)  by  themselves.  These 
have  generally  been  considered  as  very  nearly  allied  to  the  tenrecs 
(Centetidce)  of  Madagascar,  but  Mr  O.  Thomas1  is  of  opinion 
that  the  relationship  is  not  really  very  close,  the  similarity  in  the 
structure  of  their  molars  being  merely  a  generalised  character. 
Both,  however,  probably  indicate  an  ancient  group,  which  has 

1  Proc.  Zool.  Soc.  1892,  p.  500. 


III.]  CARNIVORA.  71 

migrated  from  the  higher  latitudes  of  the  northern  hemisphere  to 
find  a  refuge  in  these  two  far  distant  localities.  It  may  be 
mentioned  that  the  Solenodontidce  and  Centetidce,  together  with  the 
Potamogalida  of  western,  and  the  Chrysochloridcz,  or  golden  moles, 
of  southern  Africa,  constitute  a  section  of  the  order  distinguished 
from  all  the  other  forms  by  having  the  cusps  on  their  upper  molar 
teeth  arranged  in  the  form  of  the  letter  V,  instead  of  in  that  of  a 
W ;  and  it  will  not  fail  to  be  noticed  that  the  whole  group  is  now 
exclusively  a  southern  one.  Whether  it  is  represented  in  the  Santa 
Cruz  beds  is  not  quite  certain,  although  one  lower  jaw  has  been 
referred  to  it  by  Dr  Ameghino1  under  the  name  of  Necrolestes. 
Be  this  as  it  may,  it  is  clear  that,  with  the  exception  of  the  afore- 
said shrews,  insectivores  with  W-shaped  molars  are  entirely  want- 
ing in  the  realm ;  and  that  such  V-shaped  types  as  still  exist,  or 
formerly  occurred,  evidently  indicate  an  ancient  northern  group. 
Other  instances  of  the  survival  in  South  America  and  Madagascar 
of  allied  forms  will  be  noticed  in  the  sequel,  and  admit  of  a 
somewhat  similar  explanation. 

Although  fairly  well  represented  at  the  present  day,  the  carni- 
vores of  this  realm  have  but  few  absolutely  charac- 

.  Carnivora. 

tenstic  forms,  and  as  no  remains  of  the  true  Carni- 
vora occur  in  the  Santa  Cruz  beds,  the  whole  of  them  may  be 
regarded  as  comparatively  late  immigrants  from  the  north.  The 
civet  family  (  Vsvtrrida)  and  hyaenas  (Hyanidcz)  are  totally  wanting 
at  all  epochs,  as  indeed  they  are  throughout  the  whole  of  the  New 
World;  but  the  weasel  tribe  (Mustelidce)  have  a  comparatively 
small  number  of  representatives.  Cats  (Felidcz)  on  the  other 
hand  are  numerous,  although  several  species,  and  more  especially 
the  puma  (Felts  concolor),  range  to  a  greater  or  less  extent  into 
North  America.  Such  a  cosmopolitan  genus  is,  however,  of  no 
importance  whatever  from  a  distributional  point  of  view;  and 
much  the  same  may  be  said  of  the  extinct  sabre-tooths  (Machar- 
odus),  which  were  distributed  in  Tertiary  times  throughout  the 
entire  northern  hemisphere.  Unknown  in  the  deposits  of  Monte 
Hermoso  and  Santa  Cruz,  this  genus  is  represented  by  a  gigantic 
species  in  the  Pampean,  which  undoubtedly  reached  the  country 

t 

1  Bol.  Ac.  Cordoba,  Vol.  xm.  p.  364  (1894). 


72  THE    NEOG^ilC   REALM.  [CHAP. 

from  the  north  in  company  with  the  other  late  immigrants.  The 
dog  tribe  (Canidcs)  likewise  comes  under  the  category  of  cosmo- 
politan groups,  and  has  numerous  South  American  species, 
although  only  one  from  the  Monte  Hermoso  beds  dates  earlier 
than  the  Pampean.  It  may  be  mentioned  that  while  true  wolves1 
are  absent  from  this  realm,  all  the  continental  living  members  of 
the  genus  Cants  found  in  it  form  a  group  by  themselves,  quite 
distinct  from  the  true  foxes  of  other  regions.  Very  remarkable  is  the 
occurrence  in  the  Pampean  of  a  large  species  (C.  moreni)  perfectly 
distinct  from  all  those  now  inhabiting  the  country,  and  presenting 
some  curious  approximations  in  the  structure  of  the  skull  to 
domestic  dogs.  Peculiar  to  the  realm  is  the  bush-dog  (Icticyori), 
of  Guiana  and  Brazil, — a  small  short-haired  and  short-legged 
species  differing  from  all  other  members  of  the  family  by  the 
small  size  and  reduced  number  of  the  molar  teeth.  Its  remains 
occur  in  the  Brazilian  caves,  but  are  unknown  from  earlier 
deposits ;  and  it  may  thus  be  regarded  as  a  comparatively  late 
immigrant  from  the  north,  which  perhaps  developed  its  special 
characters  after  its  arrival  in  the  country.  Of  the  Ursidce.  there  is 
but  a  single  existing  species  in  Neogaea,  namely  the  spectacled 
bear  ( Ursus  ornatus)  of  the  highlands  of  Chili  and  Peru ;  but 
there  occur  in  the  Pampean  formation  of  the  Argentine  remains  of 
an  allied  extinct  genus  known  as  Arctotherium,  another  species  of 
the  same  genus  being  recorded  from  the  superficial  deposits  of 
California.  Far  more  characteristic  of  the  realm  are  the  raccoons 
and  coatis  (Procyonid<z\  although  several  of  these  are  common  to 
North  America.  Till  recently  it  was  considered  that  this  family 
was  entirely  restricted  to  the  New  World,  but  the  Oriental  cat-bear 
or  panda  (Ailurus)  is  now  included ;  and  since  fossil  remains  of 
the  latter  genus  have  been  discovered  in  the  Pliocene  of  England, 
while  those  of  raccoons  and  of  the  extinct  genus  Leptarctus  occur 
in  that  of  the  United  States,  it  is  practically  certain  that  the  group 
was  formerly  widely  distributed  over  the  northern  hemisphere. 
Among  this  family  the  raccoons  (Procyon)  extend  over  most  parts 
of  both  North  and  South  America ;  the  coatis  (Nasua}  range  from 
Mexico  to  Paraguay;  the  kinkajou  (Cercoleptes)  is  found  from 

1  The  Falkland  Island  wolf  (Cants  antarcticus)  forms  a  remarkable  ex- 
ception. 


III.]  UNGULATES.  73 

Mexico  to  Peru  and  Brazil;  while  the  two  species  of  the  genus 
Bassariscus  are  inhabitants  of  the  southern  United  States,  Mexico, 
and  Central  America.  From  the  Tertiaries  of  Catamarca  and 
Parana  there  has  been  described  the  extinct  genus  Cyonasua, 
differing  from  the  c.oatis  in  the  form  of  its  teeth ;  this  genus 
showing  that  the  family  had  obtained  an  entrance  into  South 
America  as  early  as  the  Pliocene  period,  although  it  is  quite 
unknown  in  the  Santa  Cruz  epoch.  In  the  Mustelidce. — where  all 
reference  to  the  cosmopolitan  otters,  of  which  there  is  one  Brazilian 
species,  may  be  omitted — the  southern  skunks  (Conepatus],  which 
have  mostly  but  thirty-two  teeth,  are  now  practically  character- 
istic of  this  realm — although  the  common  species  ranges  into 
Texas — and  their  remains  occur  fossil  in  the  caverns  of  Brazil. 
The  true  skunks  (Mephitis),  on  the  other  hand,  which  have 
thirty-four  teeth,  are  North  American,  although  one  species 
ranges  as  far  south  as  Guatemala ;  and  since  the  whole  group  is 
unknown  in  the  earlier  Tertiary  deposits  of  the  Argentine,  it  may 
likewise  be  considered  a  recent  immigrant  from  the  north.  The 
same  is  true  of  the  genus  (Galictis)  now  represented  by  the 
South  American  grison  and  tayra,  since  remains  of  this  group  of 
mustelines  occur  in  the  Plistocene  of  the  United  States,  as  well 
as  in  the  Brazilian  caves. 

Of  far  more  importance  than  either  of  the  preceding  orders 
are  the  hoofed  mammals  or  ungulates,  since  here  we 
meet  with  certainly  three,  and  probably  four  extinct 
subordinal  groups  exclusively  confined  to  this  realm,  while  a  large 
number  of  the  existing  families  are  unrepresented  even  in  the 
Pampean  formation.  Whereas  this  order  is  entirely  absent  from  the 
West  India  Islands,  South  America  at  the  present  day  is  singularly 
poor  in  ungulates.  The  only  existing  forms  are  the  peccaries 
(Dicotyles),  which  are  peculiar  to  the  New  World;  certain  deer 
belonging  to  a  genus  ( Cariacus)  which  is  likewise  confined  to  the 
western  hemisphere,  and  a  Chilian  form  constituting  a  genus 
(Pudud]  by  itself;  the  exclusively  South  American  guanacos  and 
vicunas,  which,  together  with  their  domesticated  allies  constitute 
the  genus  Lama;  and  tapirs  (Tapirus).  The  first  three  of  these 
belonging  to  the  Artiodactyla,  while  the  last  alone  represents  the 
Perissodactyla,  or  odd-toed  division  of  the  order.  At  all  periods 


74  THE    NEOG^EIC    REALM.  [CHAP. 

of  its  history  true  pigs  (Sus),  hippopotami  (Hippopotamus],  camels 
(Came/us),  chevrotains  (Tragulida\  giraffes  (Giraffida),  true  deer 
(Cervus),  antelopes,  sheep,  goats,  and  oxen,  together  with  a 
number  of  extinct  forms  connecting  the  ruminants  with  the  pigs, 
have  been  conspicuous  for  their  absence.  The  same  is  true, 
among  the  perissodactyles,  of  the  rhinoceroses  (Rhinocerotidce) 
and  the  extinct  palseotheres  (Pal&otheriida)  and  lophiodons 
(Lophiodontida)  of  Europe,  and  the  uintatheres  {Uintatheriida) 
and  titanotheres  ( Titanotheriidce]  of  the  United  States  ;  while  the 
true  elephants  (Elephas]  among  the  Proboscidea  have  likewise 
been  always  absent. 

Of  the  existing  South  American  ungulates  the  peccaries 
belong  to  a  family  (DicotylidcB)  which  is  abundantly  represented  in 
the  Tertiary  formations  of  the  United  States  ;  while  in  those  of 
Europe  and  Asia  there  occur  allied  forms  apparently  connecting 
the  peccaries  with  the  true  pigs.  On  the  other  hand,  in  South 
America  their  remains  occur  only  in  the  superficial  and  cavern- 
deposits,  so  that  there  can  be  no  doubt  as  to  their  late  intrusion 
into  the  country  from  the  north.  The  vicunas  and  guanacos  are 
the  western  representatives  of  a  family  (Cameltda)  whose  other 
members  are  Asiatic  and  African,  and  of  which  the  past  history 
seems  to  have  been  very  similar  to  that  of  the  last  group.  Fossil 
camels  occur  in  the  Pliocene  of  India,  while  a  host  of  extinct 
genera  more  or  less  closely  allied  to  the  living  South  American 
forms  occur  in  the  Tertiaries  of  the  United  States ;  and  since  in 
Argentina  and  Brazil  remains  of  Lama  and  the  related  types 
occur  only  in  the  Monte  Hermoso,  Pampean,  and  cavern  de- 
posits, there  can  be  no  hesitation  in  regarding  the  group  as  com- 
paratively recently  immigrant  into  the  country.  The  deer  of  the 
genus  Cariacus  are  likewise  only  known  in  South  America  from 
the  Pampean  and  some  other  of  the  later  Tertiaries,  as  well  as  the 
Brazilian  caves,  while  in  the  Pliocene  of  the  United  States  they 
appear  to  be  represented  by  the  ancestral  Blastomeryx;  and  these 
accordingly  come  under  the  category  of  intruders  from  the  north. 
As  regards  the  tapirs,  the  genus  and  family  now  presents  a 
remarkable  instance  of  discontinuous  distribution,  one  species 
being  confined  to  the  Malayan  countries,  while  all  the  others  are 
South  American.  Whereas  in  the  realm  under  consideration  re- 


III.]  HORSES.  75 

mains  of  these  animals  occur  only  in  the  superficial  deposits,  they 
are  met  with  abundantly  in  the  Miocene  and  Pliocene  deposits 
of  Europe  and  Asia,  as  well  as  in  the  United  States;  and  it  is 
accordingly  clear  that  the  group  was  once  widely  distributed  over 
the  northern  hemisphere,  whence  its  surviving  members  have 
wandered  southwards  to  Malaysia  in  the  east  and  South  America 
in  the  west. 

Till  introduced  by  the  early  Spanish  settlers,  horses,  which  are 
now  so  abundant  in  the  pampas,  were  totally  un-  Horses 
known  in  South  America  in  a  living  state,  although 
their  fossilised  remains  occur  commonly  in  the  Pampean,  as  well 
as  in  the  somewhat  older  deposits  of  Parana,  and  Monte  Hermoso. 
They  are,  however,  entirely  unknown  in  the  Santa  Cruz  beds. 
Some  of  these  fossil  Argentine  horses  belong  to  the  typical  genus 
Equus\  while  others,  on  account  of  the  simpler  structure  of  their 
molar  teeth  and  the  great  length  of  the  slits  in  the  skull  beneath 
the  nasal  bones,  are  referred  to  a  separate  genus,  under  the  name 
of  Hippidium.  A  third  genus  (Onohippidimn)  is  distinguished 
from  the  latter  by  a  large  lachrymal  depression  in  the  bones  of  the 
sides  of  the  face,  corresponding  to  the  so-called  larmier  of  the  deer. 
Fossil  species  of  Equus  occur  in  the  Plistocene  deposits  of  the 
whole  northern  hemisphere,  while  those  of  the  United  States  yield 
remains  of  a  genus  (Pliohippus]  nearly  allied  to  the  South  American 
Hippidium;  and  as  both  these  genera  are  the  descendants  of 
extinct  forms  whose  remains  occur  in  the  older  Tertiaries  of  the 
same  hemisphere,  the  extinct  South  American  horses  must  likewise 
be  classed  with  the  groups  that  have  entered  the  country  from  the 
north.  Why  these  Plistocene  South  American  Equida  became 
extinct  in  a  country  so  admirably  suited  to  their  existence  as 
Argentina,  is  a  question  to  which  it  is  impossible  to  find  a 
satisfactory  answer.  With  the  horses  we  reach  the  last  of  the 
Neogaeic  representatives  of  the  more  typical  ungulates,  that  is  to 
say  the  Artiodactyla  and  Perissodactyla ;  and  we  pass  on  to  the 
other  extinct  subordinal  groups,  at  least  three  of  which,  as  already 
said,  are  peculiar  to  it. 

From  the  preceding  paragraphs  it  will  be  apparent  that,  if  we 
except  the  deer,  horses,  and  the  guanacos  and  their 
allies,  South  America,  so  far  as  the  Artiodactyla  and 


THE    NEOG^EIC   REALM. 


[CHAP. 


III.]  LITOPTERNA.  77 

Perissodactyla  are  concerned,  was  exceedingly  poorly  off  for 
ungulates  during  the  Plistocene  epoch.  Nevertheless,  the 
country  was  very  rich  in  hoofed  mammals,  not  only  during  the 
Pampean,  but  likewise  during  the  Santa  Crucian  epoch,  and  in 
this  respect  it  was  quite  as  peculiar  as  it  is  in  its  edentates.  It  is 
not  a  little  remarkable  that  three  of  the  extinct  subordinal  groups 
of  the  order  which  are  confined  to  this  realm  exhibit  a  more  or 
less  decided  approximation,  more  especially  in  the  structure  of 
their  molar  teeth,  to  the  earlier  northern  Tertiary  representatives 
of  the  Perissodactyla.  Hence  it  is  probable  that  all  the  four 
suborders  in  question  have  originated  from  a  common  ancestral 
stock,  although  apparently  before  the  perissodactyles  were  differen- 
tiated from  an  earlier  group  known  as  the  Condylarthra.  The 
date  when  the  ancestors  of  the  South  American  forms  reached 
their  present  home  is,  however,  enveloped  in  mystery,  and 
although  it  is  fairly  certain  that  such  ancestors  had  a  northern 
origin,  yet  it  is  highly  improbable  that  they  entered  South  America 
from  that  direction. 

The  first  of  the  three  extinct  subordinal  groups  in  question, 
for  which  the  name  of  Litopterna  has  been  proposed,  is  the  one 
showing  the  nearest  parallelism  with  the  Perissodactyla,  and  is 
typified  by  the  genus  Macrauchenia,  of  which  the  skeleton  is 
figured  in  the  accompanying  illustration.  In  this  group  the  cheek- 
teeth approximate  in  general  structure  to  those  of  the  well-known 
European  Oligocene  genus  Palaotherium,  although  in  the  typical 
genus  Macrauchenia  they  have  been  so  modified  as  to  render  the 
resemblance  obscure.  An  essential  characteristic  of  the  upper  molar 
teeth  (fig.  13)  is  to  be  found  in  the  presence  of  two  distinct  lobes 
to  their  outer  walls.  The  toes  of  both  fore  and  hind  feet  are 
elongated,  and  constructed  on  the  same  general  plan  as  those  of 
the  Perissodactyla,  never  exceeding  three  in  number  on  each  foot, 
and  the  middle  one  being  symmetrical  in  itself.  Moreover  the 
astragalus  of  the  tarsus  or  heel-joint  resembles  the  corresponding 
bone  of  the  latter  group  in  having  a  deep  pulley-like  groove  on  its 
upper  surface  for  the  articulation  of  the  tibia,  although  inferiorly 
it  is  unlike.  The  calcaneum,  or  heel-bone,  on  the  other  hand, 
resembles  that  of  the  Artiodactyla  in  bearing  a  small  facet  for  the 
articulation  of  the  fibula,  or  small  bone  of  the  leg.  A  more 


THE    NEOG^IC   REALM. 


[CHAP. 


FlG.   II.    SKELETON  OF  HIND  FOOT  OF  AN  EXTINCT  SPECIES  OF  RHINOCEROS. 

(To  exhibit  Perissodactyle  type  of  structure.} 


FlG.    12.      LEFT   CARPUS   AND    METACARPUS    OF    HYRAX   AND    ELEPHANT. 

(Showing  linear  and  alternating  arrangements  of  carpal  bones.} 


III.]  MACRAUCHENIA.  79 

important  difference,  from  both  the  Perissodactyla  and  Artio- 
dactyla,  occurs  in  the  structure  of  the  carpus  (wrist)  and  tarsus 
(ankle),  in  both  of  which  the  two  rows  of  small  component  bones 
are  arranged  in  vertical  series  one  above  another,  instead  of 
overlapping  and  interlocking;  this  so-called  linear  arrangement 
being  a  more  primitive  type  than  the  interlocking  or  alternating 
arrangement  characterising  the  two  existing  suborders.  The 
vertebrae  of  the  neck  are  elongated,  with  flat  terminal  faces,  and 
have  the  vertebral  artery  piercing  the  sides  of  the  neural  arch 
in  a  manner  elsewhere  found  only  in  the  camel  family  and  the 
great  anteater.  In  the  femur,  or  thigh-bone,  the  projection  known 
as  the  third  trochanter  is  much  less  developed  than  in  the  Perisso- 
dactyla. All  the  members  of  the  group  were  long-limbed,  long- 
necked,  and  slenderly  built  animals,  Macrauchenia  itself  being 
as  large  as  a  camel,  although  the  genera  from  the  Santa  Cruz 
beds  of  Patagonia  were  represented  by  species  of  much  smaller 
dimensions. 

Of  the  typical  genus  Macrauchenia  fossil  remains  occur  not 
only  in  the  Pampean  formation,  but  likewise  in  the  superficial 
deposits  of  both  Patagonia  and  Bolivia.  The  most  curious  feature 
about  this  remarkable  animal  is  the  position  in  the  skull  of  the 
aperture  for  the  nostrils,  which  instead  of  being  situated  at  the 
extremity  of  the  muzzle,  is  placed  in  the  middle  of  the  forehead, 
between  the  eyes.  Otherwise  the  skull  is  not  unlike  that  of  a 
horse  in  general  contour.  The  teeth,  which  are  forty-four  in 
number,  and  form  a  continuous  uninterrupted  series,  are  a  special- 
ised modification  of  the  type  of  those  of  the  undermentioned 
genus ;  their  crowns  being  taller,  with  a  more  complicated  ar- 
rangement of  folds.  In  nearly  all  points  of  its  structure,  especially 
in  the  number  of  the  teeth,  and  the  absence  of  large  tusks,  as  well 
as  in  the  structure  of  the  wrist-  and  ankle-joints,  Macrauchenia 
is  a  very  primitive  kind  of  animal.  In  the  Santa  Cruz  beds  of 
Patagonia  the  family  to  which  this  genus  belongs  is  represented 
by  several  smaller  animals,  such  as  those  named  Oxyodonto- 
therium,  in  which  the  aperture  for  the  nostrils  occupied  a  more 
normal  position  in  the  skull,  and  the  crowns  of  the  molar  teeth 
were  shorter  and  of  a  more  simple  structure.  The  crown-surfaces 
of  worn  upper  molar  teeth  from  the  right  side  of  the  jaw  of  both 


8o 


THE    NEOG^IC   REALM. 


[CHAP. 


genera  are  shown  in  the  accompanying  figures  (13  and  14);  the 
letters  indicating  the  corresponding  elements  in  each. 


FlG.    13.      GRINDING   SURFACE   OF    RIGHT    UPPER    MOLAR   OF   Macrauchenia. 


FlG.    14.       GRINDING    SURFACE   OF    RIGHT    UPPER    MOLAR    OF 

Oxyodontotherium . 

The  second  family,  or  Proterotheriidce,  is  confined  to  the  Monte 
Hermoso,  Parana,  and  Santa  Cruz  beds  of  Patagonia,  in  the  last 
of  which  it  is  represented  by  the  genera  Proterotherium  and  Dia- 
diaphorus.  In  these  animals,  none  of  which  much  exceeded  a 
sheep  in  size,  the  upper  molar  teeth  are  much  more  like  those  of 
the  European  Palceotherium ;  and,  in  place  of  forming  a  continuous 
series,  the  teeth  are  interrupted;  one  pair,  both  in  the  upper  and 
lower  jaw,  being  developed  into  tusks.  In  structure  the  feet  pre- 
sented a  general  resemblance  to  those  of  the  extinct  three-toed 
horses  (Hipparion)  of  the  northern  hemisphere,  but  in  some  in- 
stances the  toes  were  reduced  to  a  single  one,  thus  showing  a 


III.] 


ASTRAPOTHERIA. 


8l 


Astrapotheria. 


curious  parallelism  in  the  development  of  this  group  to  that  of  the 
horses. 

A  second  sub-order  of  ungulates,  entirely  confined  to  the  Santa 
Cruz  and  Patagonian  beds  of  Patagonia,  is  repre- 
sented by  the  two  generic  types  known  as  Astrapo- 
therium  and  Homalodontotherium,  each  of  which  constitutes  a 
family  by  itself.  Rivalling  the  rhinoceros  in  bulk,  both  of  these 
extraordinary  creatures  differ  from  the  last  group  by  the  structure 
of  their  molar  teeth,  which  approximate  to  those  of  the  Rhino- 
cerotidce,  those  of  the  upper  jaw  having  a  continuous  outer  wall,  not 
divided  into  lobes.  In  the  ankle-joint  the  astragalus  differs  from 
that  of  the  preceding  group  in  having  its  upper  surface  flat,  thus 
resembling  that  of  the  elephants.  In  both  the  wrist  and  ankle 


FlG.  15.     GRINDING   SURFACE  OF   RIGHT   UPPER   MOLAR  OF  Astrapotheiium. 

the  component  bones  were  arranged  on  the  linear  plan,  and  not 
improbably  each  foot  had  five  toes.  The  vertebrae  of  the  neck 
were  short,  with  flat  terminal  faces ;  and  in  correlation  with  this 
shortness  of  the  neck,  the  limbs  and  feet  were  more  or  less 
abbreviated.  In  the  first-named  of  the  two  genera  each  jaw  was 
provided  with  an  enormous  pair  of  tusks,  wearing  obliquely  against 
L.  6 


82  THE    NEOG^IC   REALM.  [CHAP. 

one  another  like  those  of  a  wild  boar ;  and  although  there  were 
no  small  teeth  in  the  upper  jaw,  the  lower  jaw  carried  three  pairs  of 
spatulate  incisors  quite  unlike  those  of  any  other  known  animal. 
On  the  other  hand,  Homalodontotherium  takes  its  name  from  having 
its  forty-four  teeth  arranged  in  a  continuous  even  series,  unbroken 
either  by  tusks  or  by  gaps.  Whereas  the  molar  teeth  of  Astrapother- 
ium  present  a  marked  resemblance  to  those  of  the  rhinoceroses, 
it  is  noteworthy  that  those  of  the  allied  genus  make  a  certain 
approximation  to  the  corresponding  teeth  of  an  extinct  ungulate 
from  the  European  Oligocene  known  as  Cadurcotherium,  ap- 
parently more  or  less  closely  allied  to  the  rhinoceroses.  It  is, 
however,  quite  improbable  that  it  is  to  these  allied  forms  we 
have  to  look  for  the  origin  of  the  peculiar  South  American 
ungulates,  seeing  that  they  must  have  branched  off  from  the  primi- 
tive stock  at  an  earlier  stage. 

The  third  extinct  subordinal  group,  or  Toxodontia,  takes  its 
name  from  a  gigantic  species  from  the   Pampean 

Toxodonts. 

formation  to  which  Owen,  on  account  of  the  pecu- 
liarly curved  form  of  the  molar  teeth,  gave  the  name  of  Toxodon. 
Rivalling  the  largest  existing  rhinoceros  in  point  of  size,  the 
Toxodon  has  at  first  sight  very  much  the  general  appearance  of 
one  of  those  animals,  having  a  massive  skull,  short  limbs  and 
neck,  and  three-toed  feet.  The  middle  toe  is,  however,  scarcely 
larger  than  the  lateral  ones  ;  and  while  the  bones  of  the  wrist  are 
arranged  in  the  alternating  manner,  those  of  the  tarsus  are  placed 
in  a  linear  series,  and  the  astragalus  has  a  nearly  flat,  instead  of  a 
grooved,  superior  surface ;  the  terminal  faces  of  the  vertebrae  of 
the  neck  being  also  flat,  instead  of  articulating  together  by  ball- 
and-socket  joints.  Omitting  mention  of  other  more  or  less  strongly 
pronounced  peculiarities  in  the  structure  of  the  limbs,  attention 
may  be  specially  directed  to  the  teeth,  which  differ  from  those  of 
all  existing  ungulates  in  that  the  whole  of  them  grow  continuously 
throughout  the  life  of  their  owner,  without  ever  forming  roots. 
The  front  or  incisor  teeth  are  chisel-like,  and  thus  resemble  in 
form,  although  not  in  number,  those  of  a  beaver  or  rabbit;  and, 
indeed,  in  the  general  conformation  of  the  teeth,  as  well  as  in  the 
non-development  of  roots,  the  whole  dentition  of  this  animal 
presents  a  most  curious  similarity  to  that  of  the  rodents.  Some 


III.] 


TOXODONTS. 


idea  of  the  huge  size  of  these  animals  may  be  gathered  from  the 
fact  that  the  skull  may  measure  as  much  as  a  yard  in  length. 
Whereas  Toxodon  itself  is  confined  to  the  Pampean  beds,  it  is 
represented  in  the  somewhat  older  deposits  of  Monte  Hermoso 
and  Catamarca  by  allied  forms  known  as  Toxodontotherium  and 
Xotodon. 


FIG.  1 6.     SKULL  OF  Nesodon.     Much  reduced. 

In  the  Santa  Cruz  beds  of  Patagonia  the  place  of  the  fore- 
going is  taken  by  the  allied  genus  Nesodon,  likewise  belonging  to 
the  same  family  Toxodontidce.  In  these  animals,  of  which  the 
skull  is  shown  in  the  figure,  the  molars  retain  marked  resem- 
blances to  those  of  the  rhinoceroses,  which  have  been  lost  in  the 
more  specialised  Toxodon  ;  these  teeth  growing  for  a  considerable 
period,  yet  late  in  life  developing  roots  in  the  ordinary  manner. 
The  front  teeth  are  very  peculiar,  the  canines  remaining  small 
throughout  life,  but  the  second  pair  of  incisors  in  the  upper,  and 
the  third  in  the  lower  jaw  growing  beyond  the  others  to  form  large 
tusks  in  the  adult,  and  never  developing  roots.  In  their  rooted 
molars  the  Nesodons  depart  less  widely  from  the  primitive  Peris- 

6—2 


84 


THE    NEOG^EIC   REALM. 


[CHAP. 


sodactyle  type  than  do  their  more  specialised  descendants  the 
Toxodons,  although  there  are  no  forms  known  from  other  parts  of 
the  world  to  which  they  present  any  direct  relationship  or  show  any 
marked  resemblance. 

A  second  family  ( Typotheriidce)  of  the  sub-order  is  represented 
in  the  Pampean  by  the  typical  genus  Typotherium  and  in  the 
Santa  Cruz,  or  Patagonian,  beds  by  the  allied  Trachytherus.  Both 
these  have  a  dentition  still  more  like  that  of  the  rodents ;  the 
front,  or  incisor  teeth  being  reduced  to  a  single  chisel -like  pair  in 


FlG.    17.       IMPERFECT    PALATE   OF    Typotht 


Reduced. 


each  jaw,  while  the  molars  are  narrow,  rootless,  and  of  compara- 
tively simple  structure  and  relatively  small  size.  Moreover,  in- 
stead of  hoofs,  these  highly  modified  ungulates  appear,  like  many 


III.]  PYROTHERIUM.  85 

members  of  the  rodent  order,  to  have  had  their  toes  protected 
by  nails.  Like  as  are  these  animals  to  rodents,  the  resemblance 
to  that  group  is  carried  to  a  still  greater  extent  in  certain  small 
forms  from  the  Santa  Cruz  beds  which  have  been  named  Pachy- 
ruchus  and  Hegetot her  turn  ;  these  constituting  a  third  family, — the 
Pachyruchida.  From  the  features  referred  to,  the  reader  might 
be  disposed  to  consider  that  there  is  some  direct  genetic  affinity 
between  rodents  and  the  group  under  consideration,  but  this  is 
clearly  not  the  case,  such  resemblances  as  exist  being  solely  the 
result  of  that  parallelism  in  development  which  appears  to  have 
been  such  an  important  factor  in  the  evolution  of  mammals. 

It  may  be  well  to  mention  here  that  in  a  recent  paper1  Dr 
Noack  suggests  an  intimate  affinity  between  the  toxodonts  and 
the  existing  hyraces  (Hyracoidea)  of  Africa  and  Syria;  adding 
that  the  presumed  affinity  affords  evidence  of  a  land-connection 
between  Africa  and  South  America.  The  two  groups  are,  however, 
essentially  different  in  regard  to  the  structure  of  the  fore-foot,  in 
which  the  bones  of  the  carpus  or  wrist  are  arranged,  as  already 
shown,  in  the  alternating  manner  among  the  toxodonts,  whereas 
in  the  Hyracoidea  they  form  a  linear  series.  Moreover,  the  molar 
teeth  of  the  two  groups  are  markedly  distinct,  although  in  both 
they  approximate  to  the  perissodactyle  type.  Still  there  is  a  pos- 
sibility that  the  Hyracoidea  may  represent  a  less  specialised  branch 
which  has  originated  from  the  primitive  toxodont  stock,  but  has 
retained  the  linear  type  of  carpus ;  and  if  this  really  be  the  case, 
it  would  afford  very  strong  confirmation  of  the  view  that  South 
America  received  its  earliest  ungulates  by  way  of  Africa  and  the 
Antarctic  Continent. 

A  remarkable  ungulate  from  the  Patagonian  beds,  with  molar 
teeth  very  like  those  of  the  extinct  European  genus 
Dinotherium,  and  bearing  a  pair  of  large  tusks  in 
the  lower  jaw  at  least,  has  been  tentatively  assigned  by  the  present 
writer  to  the  Proboscidea ;  but  judging  from  the  distinction  of  the 
other  ungulates  of  the  older  Argentine  Tertiaries  from  those  of 
the  rest  of  the  world,  it  is  more  probable  that  it  represents  a 
sub-order  by  itself.     Pyrotherium,  as  the  animal  is  called,  was 

1  ZooLJahrb.-AbtheilfurSystemat.  vol.  vil.  pp.  540 — 542. 


86  THE    NEOG^EIC   REALM.  [CHAP. 

associated  with  nesodons,  astrapotheres,  and  homalodontotheres, 
which  have  been  assigned  by  Dr  Ameghino  to  genera  distinct 
from  those  of  the  Santa  Cruz  beds.  But  from  my  own  observations 
on  a  series  of  remains  of  these  animals  in  the  La  Plata  Museum 
obtained  in  association  with  those  of  Pyrotherium,  they  appear  to 
be  genetically  inseparable  from  their  Santa  Crucian  represen- 
tatives. 

Now   represented    only    by   the    living    Indian   and    African 
elephants   the   Proboscidea  were  formerly  a  some- 
dean°sb°SC1  what  extensive  subordinal  group  of  the  ungulates, 

easily  characterised  by  their  peculiar  molar  teeth 
and  tusks  (the  latter  of  which  may  be  present  either  in  the  upper 
or  lower  jaw  alone,  or  in  both) ;  their  five-toed  feet,  in  which  the 
bones  of  the  ankle  and  wrist  are  arranged  on  the  linear  plan,  and 
the  astragalus  has  a  flat  upper  surface;  and  the  presence  of  a 
trunk.  While  true  elephants  (Elephas)  are  totally  unknown  in 
South  America,  the  genus  Mastodon,  distinguished  by  the  low 
crowns  and  simpler  structure  of  the  molar  teeth,  is  represented  by 
two  species  in  the  Pampean  of  Buenos  Aires,  and  is  also  stated  to 
occur  in  the  Monte  Hermoso  beds ;  but  (assuming  the  distinct- 
ness of  Pyrotheriuni)  the  sub-order  is  unknown  in  the  older  deposits. 
Although  at  the  present  day  Neogaea  contains  no  existing  fami- 
lies of  either  carnivores  or  ungulates  absolutely  pecu- 

Rodents.  ..  .......  3  * 

liar  to  it,  the  case  is  widely  different  with  regard  to 
the  rodents,  or  gnawing  mammals.  Existing  rodents  are  divided 
into  seventeen  families,  arranged  under  four  sectional  groups ; 
nine  out  of  these  seventeen  families  occurring  in  the  Neogaeic 
realm,  among  which  four  are  absolutely  peculiar  to  it.  A  fifth 
(Octodontida)  is  mainly  South  American  and  West  Indian  although 
possessing  a  few  representatives  in  Africa  south  of  the  Sahara,  and 
a  sixth  (Hystricida)  has  two  genera  which  are  practically  only 
South  American.  The  significance  of  these  facts  will  be  apparent 
when  it  is  stated  that  of  the  other  zoological  regions  of  the  globe 
only  two  have  any  families  of  the  order  peculiar  to  them,  and 
neither  of  these  has  more  than  two  such  families.  In  the  Ethio- 
pian region,  for  instance,  the  only  peculiar  family  is  that  of  the 
African  flying-squirrels  (Anomalurid(E\  represented  by  two  genera ; 
while  the  western  division  of  the  Holarctic  region  has  the  sewel- 


III.]  RODENTS.  87 

lels  (Haplodontida),  with  one  genus ;  and  the  Sonoran  region  the 
pouched  rats  (Geomyidce)  with  two.  It  is  further  highly  significant 
that  all  of  these  families — which  are  mainly  or  chiefly  Neotropical — 
belong  to  one  division  of  the  order,  the  Hystricomorpha,  and 
that  certain  of  them  are  represented  either  in  the  Santa  Cruz  beds 
of  Patagonia,  or  the  Parana  beds,  where  the  whole  of  the  other 
sections  and  families  are  totally  unknown.  This,  however,  is  by 
no  means  all,  for  throughout  the  Tertiaries  of  North  America 
below  the  Pliocene  there  are  no  Hystricomorpha  at  all,  and  at  the 
present  day  there  is  but  a  single  species — the  Canadian  porcupine 
(Erethizott) — inhabiting  the  whole  of  that  country.  On  the  other 
hand,  both  at  the  present  day  and  in  the  Tertiaries,  North  America 
abounds  in  Sciuromorpha  and  Myomorpha.  These  facts  clearly 
point  to  the  existence  of  an  impassable  barrier  between  North 
and  South  America  during  a  large  portion  of  the  Tertiary 
period.  Moreover,  even  at  the  present  day  the  Sciuromorpha 
are  scarcely  represented  at  all  in  Neogaea,  while  the  Myomorpha 
are  not  very  numerous. 

Of  the  squirrel-like  rodents,  constituting  the  section  Sciuro- 
morpha, and  including  the  four  existing  families  of  the  African 
flying-squirrels  (Anomalurid(z\  the  squirrels  and  marmots  (Sciur- 
idcK],  the  sewellels  (Haplodontida),  and  the  beavers  (Castoridce), 
as  well  as  the  extinct  American  Castoroidida,  the  only  living 
Neogaeic  representatives  are  certain  species  of  squirrels,  none  x>f 
which  range  south  of  Paraguay.  The  extinct  Castoroididcz  include 
large  beaver-like  rodents  with  complex  molars  like  those  of  the 
viscacha,  and  are  represented  by  the  typical  Castoroides  from  the 
Plistocene  of  the  United  States,  and  likewise  by  Amblyrhiza 
(Loxomylus)  from  caves  in  the  West  Indies,  and,  it  is  said,  also 
from  the  later  Tertiaries  of  Argentina.  This  family  is  accordingly 
a  northern  type. 

The  second  or  murine  group  of  rodents  (Myomorpha)  contains 
five  families,  namely  the  dormice  (Myoxidce),  the  jumping-mice 
and  jerboas  (Dipodidce),  the  mice  and  rats  (Muridce),  the  mole-rats 
(Spalarida),  and  the  American  pouched  rats  (Geomytda).  Out  of 
all  these,  practically  the  only  one  represented  in  the  realm  is  the 
cosmopolitan  Muridce,  although  among  the  Geomyidce  the  genus 
Heteromys  enters  the  transitional  Mexican  sub-region.  In  the 


88  THE    NEOG^IC   REALM.  [CHAP. 

Muridce,  however,  as  in  North  America,  the  true  rats  and  mice 
(Mus)  of  the  Old  World  are  entirely  wanting,  except  through 
human  introduction ;  their  place  being  taken  by  the  white-footed 
mice  (Sttomys)  common  to  the  whole  of  the  New  World,  and 
nearly  related  to  the  European  hamsters.  Being  essentially  a 
northern  type,  they  are  certainly  recent  immigrants  into  South 
America  from  the  north ;  and  the  same  is  doubtless  true  of  certain 
genera  of  the  family  now  peculiar  to  this  realm,  such  as  the  fish- 
eating  rats  (Ichthyomys\  of  Peru  and  Venezuela,  the  groove-toothed 
mice  (Rhithrodori),  one  of  which  ranges  as  far  south  as  Tierra  del 
Fuego,  and  the  Brazilian  genus  Holochilus,  which  is  another  form 
allied  to  the  hamsters.  Although  the  cotton-rat  (Sigmodon)  occurs 
in  Central  America,  and  occasionally  wanders  still  further  south, 
the  whole  tribe  of  voles  and  their  kindred  are  absent  from  the 
entire  realm. 

Represented  by  six  families,  all  of  which  occur  within  its  limits, 
the  porcupine-like  group,  or  Hystricomorpha,  may  be  regarded  as 
the  characteristic  rodents  of  Neogsea.  From  both  the  preceding 
sections  of  the  order  the  members  of  this  section  may  be  readily 
distinguished  by  the  angle  (or  hinder  inferior  projection)  of  the 
lower  jaw  taking  its  origin  from  a  prominent  ridge  running  along 
the  side  of  the  jaw,  instead  of  from  the  inferior  edge  of  the  socket 
for  the  incisor  teeth.  Of  the  families  confined  to  this  realm,  that 
of  the  cavies  (Caviidcz],  includes  heavily-built  rodents,  with  four 
front  and  three  hind  toes,  rudimental  or  short  tails,  and  the 
cheek-teeth  divided  by  transverse  folds  of  enamel  into  a  number 
of  thin  parallel  plates.  The  genera  of  this  family  include  the  true 
cavies  (Cavia] — so  well  known  through  the  domestic  guinea-pig — 
all  of  which  are  small  short-limbed  forms ;  the  larger  and  taller 
Patagonian  cavy  (Dolichotis};  and  the  carpincho,  or  capivara 
(Hydrocha>rus\  which  is  the  largest  living  member  of  the  order, 
and  is  characterised  by  the  large  number  of  plates  going  to  form 
the  last  molar  tooth  in  each  jaw.  Although  they  do  not  appear  to 
have  been  recorded  from  the  Santa  Cruz  beds,  remains  of  members 
of  this  family  occur  in  the  Parana1  horizon,  and  also  in  that  of 

1  There  is  some  confusion  with  regard  to  the  age  of  the  Parana  beds.  They 
are  overlain  by  marine  strata  which  have  been  identified  with  the  Patago- 
nian ;  but  it  is  more  probable  that  they  are  newer  than  the  Santa  Cruz  beds, 


III.]  RODENTS.  89 

Monte  Hermoso.  Of  the  former  Plexochairus  differs  from  Hydro- 
chorus  only  by  the  somewhat  simpler  structure  of  the  last  molar, 
Hydrochcerus  itself  occurring  in  the  Monte  Hermoso  beds.  Other 
forms  from  the  Parana  stage  are  Eucardiodon  and  Cardiotherium, 
apparently  more  nearly  allied  to  Cavia.  Here  it  should  be 
mentioned  that  certain  European  Oligocene  genera  (Issiodoromys 
and  Nesocerodori)  have  the  molars  so  like  those  of  the  cavies  that 
they  are  regarded  by  Dr  Schlosser  as  nearly  allied'  to  the  family. 
By  Prof.  Zittel  they  are,  however,  included  in  the  extinct  family 
Theridomyidce,  which  is  classed  with  the  dormice  and  certain 
other  families  in  a  group  regarded  as  intermediate  between  the 
Sciuromorpha  and  Hystricomorpha.  If,  as  would  seem  probable, 
they  are  really  allied  to  the  latter,  they  are  of  the  utmost  import- 
ance as  indicating  a  connection  between  the  middle  Tertiary 
rodent  fauna  of  Europe  with  that  of  South  America1. 

Nearly  allied  to  the  cavies  are  the  agutis  (Dasyprocta)  and 
pacas  (C&logenys),  collectively  constituting  the  family  Dasy- 
proctidcE,  and  differing  from  the  former  in  that  the  folds  of  enamel 
merely  form  notches  on  the  sides  of  the  crowns  of  the  cheek-teeth. 
In  a  fossil  state  the  family  seems  to  be  only  known  by  remains  of 
the  existing  genera  from  the  Brazilian  caves.  The  third  peculiar 
family  (Dinomyidce)  is  known  merely  by  a  single  specimen  from 
Peru.  The  only  other  family  exclusively  confined  to  the  realm  is 
that  of  the  Lagostomatidcz  (Chinchillidce],  which  includes  not  only 
the  true  chinchillas  (Eriomys)  and  Cuvier's  chinchilla  (Lagidium), 
but  likewise  the  viscacha  (Lagostomus]  of  the  Argentine  pampas. 
All  the  members  of  this  family  have  long  bushy  tails,  elongated 
hind  limbs,  and  the  cheek-teeth  divided  by  complete  transverse 
folds  of  enamel  into  thin  plates.  Lagostomus  occurs  fossil  not 
only  in  the  Pampean,  but  likewise  in  the  older  Tertiaries  of 
Parana;  while  it  may  be  doubted  whether  Pliolagostomus  and 
Prolagostomus  of  the  Santa  Cruz  beds  are  really  entitled  to  generic 
distinction.  Other  allied  forms  from  the  latter  deposits  are 

although  their  lowest  portion  is  older  than  the  Monte  Hermoso  stage.  (See 
Ameghino,  Bull.  Ac.  Cordoba,  Vol.  xiil.  pp.  260,  261,  1894.)  They  may  be 
partly  made  up  of  the  remains  of  pre-existing  beds. 

1  Dr  Schlosser  writes  me  to  the  effect  that  he  is  fully  assured  these  forms 
are  the  ancestors  of  the  Caviidce. 


90  THE    NEOG-^IC   REALM.  [CHAP. 

described  under  the  names  of  Perimys  and  Sphodromys,  and  thus 
indicate  that  the  family  is  essentially  South  American.  The 
largest  known  rodent  is  the  extinct  Megamys,  from  the  Parana 
beds ;  the  typical  species  of  the  genus  being  described  as  equal  in 
size  to  an  ox.  The  porcupines  (Hystricida),  which  form  a 
practically  cosmopolitan  family,  sufficiently  distinguished  by  their 
spiny  covering,  are  represented  in  South  America  by  the  two 
arboreal  genera  Synetheres  and  Chcztomys,  differing  from  all  their 
allies  by  their  prehensile  tails,  although  otherwise  related  to  the 
North  American  Erethizon,  with  which  they  form  a  separate  sub- 
family. Of  Chcetomys  an  extinct  representative  occurs  in  the 
cavern-deposits  of  Brazil;  and  in  the  Santa  Cruz  beds  the  family 
is  represented  by  apparently  extinct  generic  types  described  under 
the  names  of  Stiromys,  Acaremys,  and  Sciamys.  Since  fossil 
remains  of  Hystrix  date  from  the  European  Miocene  or  Oligocene, 
there  is  distinct  evidence  of  a  connection  between  the  early  South 
American  rodents  and  those  of  the  Old  World ;  while  as  Erethi- 
zon is  first  known  from  the  Plistocene  of  North  America,  it  may 
probably  be  regarded  as  a  late  immigrant  into  that  country  from 
the  south. 

The  largest  of  all  the  families  of  the  Hystricomorpha  is  that  of 
the  Octodontidal,  in  which  out  of  a  total  of  nineteen  genera 
upwards  of  fifteen  belong  to  the  realm  under  consideration,  while 
the  remaining  four  are  African  and  mainly  Ethiopian.  In  addition 
to  other  features,  the  rodents  of  this  family  have  the  crowns  of 
the  cheek-teeth  marked  by  infoldings  of  enamel  from  both  sides; 
there  are  usually  five  toes  to  each  foot ;  and  the  general  form  is 
more  or  less  rat-like.  Of  the  Neogaeic  forms,  the  typical  genus 
Octodon  is  represented  by  the  degu  of  Chili  and  Peru,  which  is  a 
large  rat -like  animal,  with  a  brush-tipped  tail;  other  species 
occurring  in  Bolivia.  The  latter  country  is  likewise  the  home  of 
the  allied  genus  Habrocoma,  the  members  of  which  vie  with  the 
chinchillas  in  the  delicate  softness  of  their  fur.  Nearly  related  are 
the  burrowing  South  American  tuco-tucos  (Ctenomys\  characterised 

1  By  some  the  family  is  divided  into  three;  viz.  Capromyidce,  with  the 
West  Indian  Capromys,  the  S.  American  Myopotamus,  and.  the  African 
Triaulacodus ;  Ctenodactylidcz,  including  the  remaining  African  forms;  and 
Octodontidce,  comprising  all  the  other  American  types. 


III.]  RODENTS.  91 

by  the  comb-like  bristles  on  the  hind  feet,  and  their  bell-like  cry ; 
two  Chilian  species,  constituting  the  genus  Spalacopus,  being 
distinguished  by  their  rudimental  ears.  Schizodon,  on  the  other 
hand,  of  which  there  is  a  single  species  from  the  Southern  Andes, 
has  the  ears  larger  than  in  the  tuco-tucos.  The  South  African 
Petromys  has  been  regarded  as  very  closely  allied  to  Spalacopus^ 
but  it  is  more  probable  it  should  be  classed  with  the  other  two 
African  genera  Ctenodactylus  and  Pectinator,  to  constitute  a 
separate  sub-family.  The  third  sub-family  includes  all  the  other 
genera,  one  of  which  (Triaulacodus1},  as  represented  by  the  cane- 
rats,  is  Ethiopian,  while  the  whole  of  the  remainder  are  Neogaeic. 
Many  of  the  species  are  of  large  size,  some  being  arboreal  and 
others  aquatic.  Of  the  latter  the  best  known  and  largest  is  the 
coypu  (Myopotamus),  widely  spread  over  South  America  ;  while 
in  the  West  Indies  the  group  is  represented  by  the  equally  large 
arboreal  hutias  (Capromys  and  Plagiodori).  The  other  seven 
genera  are  South  American,  and  include  smaller  rat-like  forms, 
which  in  the  case  of  the  two  genera  Loncheres  and  Echinomys  have 
flattened  spines  mingled  with  the  fur;  the  others  being  known 
as  Mesomys,  Dactylomys,  Cannabateomys,  Cercomys,  and  Cartero- 
don.  Several  of  the  existing  genera  occur  fossil  in  the  caves  of 
Brazil  and  the  Pampean ;  Myopotamus  also  occurring  in  the 
infra-pampean  beds  of  Parana,  together  with  the  reputedly 
extinct  forms  described  as  Orthomys  and  Morenia.  Other  extinct 
genera,  such  as  Neoremys,  Scleromys,  and  Addphomys  occur  in  the 
Santa  Cruz  deposits,  and  appear  to  be  very  closely  allied  to 
Myopotamus.  It  is  noteworthy  that  an  extinct  Octodont  (Pelle- 
grinia)  allied  to  Ctenodactylus  occurs  in  the  Plistocene  or  Pliocene 
of  Sicily;  while  Ruscinomys  from  the  Pliocene  of  France  is 
believed  to  belong  to  the  same  group.  Finally,  the  genus 
Eocardia,  together  with  certain  other  allied  forms  from  the  Santa 
Cruz  beds,  are  regarded  as  indicating  a  separate  family  (Eocar- 
ditdce)  of  the  section. 

With  regard  to  the  extinct  family  Theridomyida  from  the 
middle  and  upper  Oligocene  of  Europe,  which  includes  not  only 
Theridomys  and  Archceomys,  but  probably  also  the  above-men- 

1  The  name  Aulacodus  being  preoccupied,  that  of  Triaulacodus  is  proposed 
in  substitution. 


92  THE    NEOG^IC   REALM.  [CHAP.  III. 

tioned  Nesocerodon  and  Issiodoromys  (p.  89),  it  seems  highly 
probable  that  these  are  really  the  ancestral  forms  of  the  modern 
Hystricomorpha,  although  their  lower  jaws  approximate  to  the 
general  type  of  those  of  the  more  generalised  Sciuromorpha  and 
Myomorpha. 

The  last  section  of  the  order  (Lagomorpha),  which  includes  the 
hares  and  picas,  and  is  essentially  a  northern  one,  is  but  poorly 
represented  in  Neogaea ;  the  picas  (Lagomyidcz)  being  unknown 
there,  while  in  the  whole  of  the  realm  there  are  only  two  species  of 
Leporidce,  one  of  which  is  Brazilian. 

It  has  been  usual  in  zoological  systems  to  include  under  the 
title  of  Edentata  not  only  the  armadillos,  anteaters, 
and  sloths  of  South  America,  but  likewise  the  Old 
World  pangolins  (Manida)  and  aard-varks  (Orycteropodidaz). 
There  can,  however,  be  no  doubt  that  there  is  little  or  no 
connection  between  the  two  groups,  and  the  latter  may  accord- 
ingly be  separated  as  a  distinct  order  under  the  title  of  ErTodientia. 
In  this  restricted  sense  the  edentates  at  the  present  day  are,  per- 
haps, the  most  characteristic  and  remarkable  of  all  the  Neogaeic 
mammals.  Whereas,  however,  the  sloths  and  anteaters  are 
entirely  Neogseic,  a  few  of  the  armadillos  have  wandered  at  a 
comparatively  modern  date  as  far  north  as  Texas ;  but  this  does 
not  detract  from  their  essentially  southern  character,  seeing  that 
they  are  well  represented  in  the  Santa  Cruz  beds,  and,  if  we 
exclude  certain  remains  of  doubtful  affinity  from  the  Oligocene 
of  France,  are  quite  unknown  elsewhere.  This,  however,  is  by 
no  means  all,  since  there  are  two  extinct  families  of  the  order, 
dating  from  the  Santa  Cruz  beds,  which  were  extremely  abundant 
during  the  Pliocene  and  Plistocene  epochs ;  some  few  of  these 
having  managed  to  obtain  an  entrance  into  North  America 
about  the  Miocene  epoch.  Central  and  South  America  may 
accordingly  be  considered  as  essentially  the  home  of  the  edentates ; 
and  are  thus  broadly  demarcated  from  all  other  parts  of  the  world. 
It  would  be  superfluous  to  point  out  all  the  distinctive  features  of 
the  order,  but  it  may  be  mentioned  that  none  of  the  living  forms 
have  teeth  in  the  front  of  the  jaws ;  while  in  all  those  genera  in 
which  teeth  are  present,  these  are  of  comparatively  simple 
structure,  being  unprovided  with  a  coating  of  enamel,  growing 


94  THE    NEOG^IC   REALM.  [CHAP.III. 

continuously  without  ever  forming  roots,  and  being  mostly  very 
similar  throughout  the  series. 

The  mailed,  or  loricate  edentates  are  represented  by  the  two 
families  of  the  armadillos  (Dasypodidce]  and  glypto-  Armadillos 
donts  (Glyptodontida],  the  latter  of  which  died  out  and  Glypto  - 
at  the  close  of  the  Plistocene  or  the  commence- 
ment of  the  Recent  epoch,  whereas  the  former  is  still  abundant. 
With  the  exception  of  the  two  species  of  pichiciagos  (Chlamydo- 
phorus]  from  Mendoza  and  Bolivia,  in  which  a  solid  coat  of  mail 
is  confined  to  the  hinder  region  of  the  body,  the  members  of  both 
families  have  their  bodies  protected  by  a  bony  armour,  while 
their  heads  are  guarded  above  by  a  shield  of  the  same  nature, 
and  their  tails  are  enclosed  in  a  tubular  sheath.  Covered  exter- 
nally with  horny  shields,  like  the  shell  of  a  tortoise,  the  carapaces 
of  these  animals  are  formed  of  a  number  of  small  plates  of  bone, 
either  united  everywhere  by  their  edges  into  a  continuous  solid 
armour,  or  in  the  middle  region  of  the  body  overlapping  one 
another  like  the  tiles  on  a  roof.  In  the  true  armadillos  (of  which 
the  living  Argentine  forms  are  all  comparatively  small  creatures, 
although  one  Brazilian  species  reaches  nearly  a  yard  in  length)  the 
carapace  consists  of  a  nearly  solid  buckler  in  front  and  behind,  while 
between  the  two  are  situated  a  variable  number  of  movable  over- 
lapping bands,  which  in  some  instances  admit  of  the  body  being 
rolled  up  into  the  form  of  a  ball.  They  have  all  long  snouts,  and 
simple,  subcylindrical  teeth.  The  true  existing  armadillos  may 
be  divided  into  the  genera  Dasypus,  Xenurus,  Priodon,  Tolypeutes, 
and  Tatusia.  The  first  of  these,  in  which  there  are  six  or  seven 
movable  bands  to  the  carapace,  is  found  throughout  the  Argentine 
Tertiaries  to  the  Santa  Cruz  beds,  one  of  the  fossil  species  from 
the  higher  beds  of  the  series  having  a  skull  of  nearly  a  foot  in 
length,  and  thus  vastly  exceeding  all  its  living  congeners  in  size. 
Tatusia,  in  which  the  carapace  has  from  seven  to  nine  movable 
bands,  does  not  appear  to  be  known  below  the  Pampean  beds, 
where  it  is  represented  by  the  large  species  of  which  the 
external  skeleton  is  shown  in  the  figure  on  p.  93.  A  third  genus, 
Eutatus,  which  likewise  comprises  species  of  large  size,  and 
ranges  from  the  Pampean  to  the  Santa  Cruz  beds,  is  distinguished 
by  having  over  thirty  movable  bands  in  the  carapace.  More 


96  THE   NEOG^IC   REALM.  [CHAP. 

remarkable  than  any  is  the  extinct  Peltephilus  of  the  Santa  Cruz 
deposits,  in  which  the  teeth  form  a  continuous  series  up  to  the 
front  of  the  jaws,  while  the  skull  has  a  very  broad  snout,  and  the 
humerus  is  of  such  a  remarkable  shape  that  it  has  been  described 
as  that  of  a  monotreme.  Indeed  this  genus  seems  to  suggest 
that  edentates  are  derived  from  animals  with  a  fully  developed 
series  of  teeth  in  the  front  of  the  jaws.  The  pichiciagos  (Chlamy- 
dophorus],  which  are  unknown  before  the  Plistocene,  form  a 
sub-family  by  themselves ;  and  yet  another  sub-family  group  is 
indicated  by  the  gigantic  Chlamydotherium,  of  the  Brazilian  caves 
and  the  Pampean,  which  rivalled  the  largest  glyptodonts  in  size, 
and  had  teeth  of  a  more  complex  type  than  the  true  armadillos. 
Other  species  occur  in  the  Catamarca  and  Monte  Hermoso  Ter- 
tiaries,  although  the  genus  is  unknown  in  those  of  Patagonia. 

From  the  armadillos  and  their  immediate  kin  the  extinct 
glyptodonts  differ  in  having  the  carapace  in  the  form  of  a  con- 
tinuous solid  shield,  without  any  movable  bands  in  the  middle 
region ;  in  addition  to  which  the  skull  is  characterised  by  its 
depth  and  shortness,  while  the  teeth  form  long  fluted  prisms. 
The  internal  skeleton,  as  shown  in  figure  19,  is  characterised  by 
the  union  of  nearly  the  whole  of  the  vertebrae  of  the  back  into 
a  solid  girder  for  the  support  of  the  massive  carapace ;  and  the 
feet  are  furnished  with  much  shorter  claws  than  those  of  the 
burrowing  armadillos,  the  hinder  ones  being  almost  nail-like  in 
form.  Most  of  the  species  from  the  Pampean  formation  are 
animals  of  gigantic  dimensions,  the  length  of  the  carapace  being 
not  unfrequently  from  six  to  eight  feet ;  and  they  are  unquestion- 
ably some  of  the  most  extraordinary  creatures  that  ever  trod  the 
earth.  Although  the  majority  are  South  American,  some  members 
of  the  genus  wandered  as  far  north  as  Texas,  while  from  the 
upper  division  of  the  Loup  Fork  beds,  corresponding  to  the 
lowest  Pliocene,  a  North  American  form  has  been  described  under 
the  name  of  Carioderma. 

In  the  typical  genus  Glyptodon,  which  ranges  from  the  sand- 
dunes  and  Pampean  formation  to  the  Monte  Hermoso  beds,  the 
tail-sheath,  as  shown  in  the  annexed  figure,  is  composed  of  a 
number  of  spiny  rings,  gradually  decreasing  in  size  from  the  root 
to  the  tip,  and  the  polygonal  plates  of  the  carapace  are  each 


I 


Cs.        O 

0 


98  THE    NEOG/EIC   REALM.  [CHAP. 

ornamented  with  a  distinct  rosette-like  sculpture.  The  allied 
genus  Plohophorus,  of  which  the  remains  occur  alike  in  the 
Brazilian  caves  and  the  Catamarca  and  Monte  Hermoso  beds  of 
Argentina,  while  agreeing  with  the  last  in  the  characters  of  the 
skull,  has  a  carapace  more  like  that  of  the  undermentioned 
Panochthus,  and  a  tail-sheath  resembling  that  of  the  next  genus. 
In  addition  to  well-marked  distinctive  features  of  the  skull,  Loma- 
phorus  is  characterised  by  the  great  elongation  and  slender  form 
of  the  carapace,  which  is  produced  on  either  side  of  the  neck  in 
the  same  manner  as  in  the  armadillos,  while  its  margins  lack  the 
large  bosses  exhibited  by  the  typical  genus.  The  tail-sheath 
consists  of  a  small  number  of  rings  at  the  base,  followed  by  a 
long  terminal  tube  ornamented  with  smooth,  oval,  bony  plates, 
of  which  those  along  the  sides  and  at  the  tip  are  larger  than  the 
rest.  The  genus,  of  which  the  species  are  much  inferior  in  point 
of  size  to  those  of  Glyptodon,  has  the  same  geological  range  as 
Plohophorus. 

Another  type  of  the  family  is  represented  by  the  animals 
constituting  the  genus  Panochthus,  in  which  the  hexagonal  bony 
plates  of  the  carapace  are  arranged  in  more  distinct  rows  on  the 
sides  of  the  body ;  those  of  the  back  being  ornamented  either 
with  a  number  of  small  granular  tubercles  (as  in  the  species  here 
figured),  or  with  one  circular  central  disc  surrounded  with  several 
rows  of  much  smaller  discs.  A  more  striking  difference  is  dis- 
played in  the  structure  of  the  sheath  of  the  tail,  which  consists  at 
the  base  of  six  or  seven  large  smooth  rings  diminishing  very 
rapidly  in  diameter,  and  terminates  in  a  long  and  massive  de- 
pressed tube,  the  sides  of  which  are  ornamented  with  large 
roughened  bosses,  probably  surmounted  during  life  with  horny 
knobs,  while  the  intervening  spaces  are  covered  with  small  bony 
ossicles.  The  species  are  of  very  large  or  medium  size,  and  range 
from  the  Pampean  to  the  Monte  Hermoso  beds  in  Argentina, 
while  some  occur  in  the  Brazilian  caves.  Still  more  extraordinary 
is  the  gigantic  Dadicurus  of  the  Pampean,  represented  by  a  some- 
what smaller  form  from  the  Monte  Hermoso  beds.  Having  a 
total  length  in  a  straight  line  of  close  upon  twelve  feet,  five  of 
which  are  taken  up  by  the  ponderous  tail,  the  Pampean  repre- 
sentative of  this  genus  has  the  outer  surface  of  the  plates  of  the 


III.] 


GLYPTODONTS. 


99 


7—2 


100  THE    NEOG^IC   REALM.  [CHAP. 

carapace  smooth ;  each  being  perforated  by  three  or  four  large 
holes  for  the  passage  of  blood-vessels,  and  the  whole  being  pro- 
bably invested  with  a  continuous  leathery  skin,  instead  of  each 
disc  bearing  a  separate  horny  shield.  Commencing  with  a  small 
series  of  enormous,  narrow,  hoop-like  rings,  the  tail-sheath  termi- 
nates in  a  long,  massive,  depressed,  and  nearly  smooth  club- 
shaped  tube,  at  the  extremity  of  which  are  a  number  of  rough 
disc-like  surfaces,  apparently  for  the  attachment  of  large  horns. 

None  of  the  preceding  forms,  which  include  all  the  largest 
members  of  the  family,  range  below  the  horizon  of  the  Monte 
Hermoso,  Catamarca,  and  Parana  beds,  but  the  group  is  also 
represented  in  the  Santa  Cruz  deposits  of  Patagonia,  although  the 
single  species  found  there  is  of  much  smaller  dimensions  than  any 
of  its  later  cousins,  the  whole  length  of  the  carapace  not  exceeding 
about  two  feet.  It  is  noteworthy  that  this  earliest  known  glypto- 
dont,  on  which  the  unwieldy  name  of  Propalczohoplophorus  has  been 
conferred,  presents  certain  indications  of  affinity  to  the  armadillos 
in  the  structure  of  its  carapace,  in  which  incipient  movable  bands 
may  be  detected  on  the  margins  of  the  middle  region.  In  this 
small  size  of  their  earliest  definitely  known  representative,  the 
glyptodonts  resemble  the  under-mentioned  ground-sloths. 

Unless  the  aforesaid  remains  from  the  Oligocene  Phosphorites 
of  France  should  prove  to  belong  to  the  group,  we  are  at  present 
totally  in  the  dark  as  to  whence  both  the  glyptodonts  and  the 
armadillos  originally  came ;  and  it  is,  indeed,  quite  probable 
that,  like  the  other  members  of  the  order,  they  may  have  origi- 
nated in  South  America  (if  not  in  an  Antarctic  continent)  from 
some  at  present  quite  unknown  mammalian  type.  How  such 
creatures,  which  seem  absolutely  unassailable,  came  to  be  extermi- 
nated, is  one  of  those  questions  which  it  appears  quite  impossible 
to  answer. 

Although  they  have  not  hitherto  been  discovered  in  a  fossil 
state,  the  sloths,  constituting  the  family  Bradypodidce, 
are  just  as  characteristic  of  Neogaea  as  the  two  pre- 
ceding groups.  Their  habits,  however,  necessarily  restricting  them 
to  the  tropical  forest-districts,  their  absence  in  a  fossil  state 1  must 

1  A  presumed  fossil  sloth  was  described  from  the  Argentine,  but  the  jaw  on 
which  it  was  founded  proves  to  belong  to  the  Megalotheriidce. 


III.]  SLOTHS   AND    ANTEATERS.  IOI 

not  be  taken  as  an  indication  that  they  did  not  exist  during  the 
Pampean  epoch,  since  their  remains  are  not  likely  to  occur  in 
the  Argentine,  although  they  might  with  more  probability  be 
looked  for  in  Brazil.  On  the  other  hand,  their  specialised  struc- 
ture makes  it  highly  probable  that  they  had  not  come  into  being 
at  the  date  of  deposition  of  the  Santa  Cruz  beds.  Of  the  dimen- 
sions of  medium-sized  monkeys,  sloths  are  characterised  by  their 
short,  rounded  heads,  and  extremely  long  limbs,  armed  with  very 
elongated  curved  claws ;  in  the  genus  Bradypus  the  latter  being 
three  in  number  on  each  foot,  but  in  Cholcepus  reduced  to  two 
in  the  fore  feet.  Their  bodies  are  coated  with  very  coarse  ragged 
hair,  and  the  tail  is  wanting.  The  teeth  are  oval  prisms,  some- 
what cupped  in  the  middle  of  their  grinding  surfaces ;  but  in 
the  last-named  of  the  two  genera  the  first  pair  in  each  jaw  are 
larger  than  the  rest,  from  which  they  are  separated  by  an  interval, 
and  form  tusks  wearing  against  one  another  to  oblique  facets. 
Usually  there  are  five  upper,  and  four  lower  teeth  on  a  side. 
The  range  of  sloths  extends  from  Mexico  throughout  the  greater 
part  of  the  forest-districts,  although  they  do  not  appear  to  reach 
as  far  south  as  Paraguay. 

Likewise  unknown  in  a  fossil  condition,  the  true  anteaters,  or 
Myrmecophagidce,  constitute  another  exclusively  Neo- 

.. '  .  .  i  •      ,  Anteaters. 

gseic  family,  with  nearly  the  same  geographical  range 
as  the  sloths,  but  represented  in  Paraguay.  So  unlike  are  these  ani- 
mals to  sloths,  that  it  is  at  first  difficult  to  believe  that  there  is  any 
close  relationship  between  the  two,  and  it  is  largely  due  to  the  evi- 
dence of  the  ground-sloths  referred  to  below  that  it  has  been  possible 
to  discover  how  close  the  connection  really  is.  In  place  of  being 
rounded  and  shortened,  the  skull  in  the  present  family  is  more  or 
less  elongated  and  slender,  with  the  jaws  entirely  devoid  of  all 
vestiges  of  teeth,  and  the  tongue  long,  cylindrical,  and  extensile. 
An  equally  striking  difference  obtains  in  regard  to  the  structure  of 
the  limbs,  the  fore  foot  of  the  great  anteater  having  five  toes,  of 
which  the  middle  one  is  much  more  powerful  than  the  others, 
while  all  except  the  fifth  are  furnished  with  strong  claws.  In 
walking,  the  outer  side  and  part  of  the  upper  surface  of  the  fore 
foot  is  applied  to  the  ground ;  but  in  the  hind  limbs  the  sole 
forms  the  support  in  the  ordinary  manner.  Whereas  sloths  are 


102  THE   NEOG^IC   REALM.  [CHAP. 

highly  specialised  as  regards  the  structure  of  their  limbs,  in  the 
present  group  the  greatest  degree  of  specialisation  shows  itself  in 
the  skull,  in  the  majority  of  the  species.  There  are  but  three 


FlG.    22.      TAMANDUA   ANTEATER. 

members  of  the  family,  each  representing  a  genus  by  itself,  namely 
the  great  anteater  (Myrmecophaga\  the  tamandua  (Tamandua],  and 
the  two-toed  or  little  anteater  ( Cycloturus] ;  the  latter  being  ex- 
clusively arboreal  in  its  habits. 

The  foregoing  remarks  on  some  of  the  structural  features  of 

the  sloths  and  anteaters  will  the  more  easily  enable 
si?th°sUnd"  the  reader  to  understand  the  peculiarities  of  the 

extinct  group  of  ground-sloths.  They  have  been 
divided  into  a  large  number  of  genera  and  several  families;  but 
the  former  may  be  considerably  reduced,  and  the  whole  of  them 
included  in  the  single  family  Megalotheriidce.  Ranging  in  the 
Argentine  from  the  Santa  Crucian  to  the  Pampean  epoch  and  the 
overlying  sand-dunes,  the  family  has  a  geographical  distributional 
area  including  North  America  as  far  as  Kentucky.  The  South 
American  forms  vastly  exceed  those  of  N.  America  in  point  of 
number;  and  whereas  the  latter  are  found  only  in  deposits  of 
upper  Pliocene  and  Plistocene  age,  the  former,  as  we  have  seen, 
extend  downwards  to  the  Miocene.  The  members  of  this  family 
may  be  denned  as  edentates  without  a  carapace,  the  skull  and 
dentition  of  the  general  type  of  those  of  the  sloths,  and  the 


III.]  GROUND-SLOTHS.  IO3 

limb-bones  and  vertebrae  like  those  of  the  anteaters.  The  skull  is, 
however,  somewhat  more  elongate  than  in  the  former,  and  in  the 
case  of  the  genus  Scelidotherium  approximates  to  that  of  the  latter. 
The  Plistocene  forms  include  by  far  the  largest  representatives  of 
the  order,  the  Megalotherium1  attaining  a  total  length  of  about 
eighteen  feet,  with  a  bodily  bulk  fully  as  great  as  that  of  an 
elephant.  Whereas  all  the  members  of  the  family  whose  remains 
occur  in  the  Plistocene  walked  on  the  outer  sides  of  their  feet,  in 
the  small  ancestral  Patagonian  forms  this  specialised  character 
seems  to  have  been  less  developed. 

The  typical  genus  Megalotherium — which  includes  several 
species,  ranging  in  time  from  the  Monte  Hermoso  and  Cordoba 
beds  to  the  Pampean,  and  in  space  from  Argentina  and  Chili  to 
South  Carolina  and  Texas — is  sufficiently  distinguished  by  having 
the  teeth  in  the  form  of  large  quadrangular  prisms,  sometimes 
measuring  as  much  as  a  foot  in  length,  and  wearing  on  their  sum- 
mits into  a  pair  of  transverse  ridges,  owing  to  the  presence  of  layers 
of  unequal  hardness.  The  allied  genus  Mylodon,  including  smaller 
forms  which  may  be  compared  in  size  to  rhinoceroses,  differs  from 
the  preceding  in  the  structure  of  the  teeth,  which  are  similar  to 
those  of  the  sloths ;  the  skull,  as  shown  in  the  figure  on 
p.  104,  being  comparatively  short,  with  the  teeth  extending  nearly 
up  to  the  extremities  of  the  jaws.  In  the  skeleton  of  this  genus 
the  limbs  are  of  moderate  length  and  very  powerful.  The  two 
outer  toes  of  the  fore  feet  are  rudimental  and  clawless,  but  the 
three  innermost  provided  with  claws,  of  which  the  third  is  much 
larger  than  either  of  the  others,  this  discrepancy  being  carried 
to  a  still  greater  extent  in  Megalotherium.  It  will  be  observed 
that  the  creature  walked  on  the  outer  sides  and  part  of  the  upper 
surfaces  of  the  fore  feet  after  the  manner  of  a  sloth ;  but,  unlike 
the  latter,  only  the  outer  sides  of  the  hind  feet  were  applied  to  the 
ground ;  the  great  middle  toe,  which  in  Megalotherium  carried  a 
gigantic  claw,  not  touching  the  ground  at  all.  In  the  structure  of 
their  feet  these  animals  are  thus  more  like  anteaters  than  sloths, 
although  the  hinder  pair  are  of  a  somewhat  more  specialised 
structure  than  in  the  latter.  It  may  be  mentioned  that  the 

1  The  name  Megatherium  clearly  requires  amendment  to  Megalotherium. 


104 


THE    NEOG^IC   REALM. 


[CHAP. 


conformation  of  their  teeth  indicates  that  the  ground-sloths  were 
vegetable-feeders,  and  it  is  probable  that  they  subsisted  largely 
upon  the  young  twigs  and  leaves  of  trees,  which  may  have  been 


FIG.  23.     UNDER-SURFACE  OF  SKULL  OF  Mylodon.     (Reduced.) 

brought  within  reach  by  the  animals  rearing  themselves  up 
against  the  trunks  and  pulling  down  the  boughs  with  their  fore 
paws.  The  present  treeless  condition  of  the  Argentine  pampas 
suggests  that  the  ground-sloths  were  grazing  rather  than  browsing 


III.]  GROUND-SLOTHS.  IO5 

animals,  but  their  structure  is  not  in  favour  of  this  view,  and  it 
must  be  remembered  that  their  remains  are  likewise  met  with  in 
Brazil,  which  was  probably  always  as  well  wooded  as  at  the  present 
day.  The  disappearance  of  forests  from  the  pampas  cannot,  in- 
deed, be  regarded  as  more  marvellous  than  the  extinction  of  its 
Plistocene  mammals.  In  the  sand-dunes  near  the  coast  at  Buenos 
Aires  bones  of  some  of  the  ground-sloths,  as  well  as  of  glyptodonts, 
have  been  found  in  association  with  human  remains,  so  that  their 
extinction  is  an  event  of  comparatively  recent  date.  The  genus  is 
typically  represented  by  Mylodon  harlani  of  the  Plistocene  of 
Kentucky  and  other  parts  of  North  America,  but  is  nevertheless 
essentially  South  American,  ranging  in  Argentina  from  the  Pam- 
pean  beds  to  those  of  Parana  and  Monte  Hermoso.  The  allied 
genus  Megalonyx  is  exclusively  restricted  to  the  North  American 
Plistocene  and  Upper  Pliocene  ;  and  here  may  be  repeated  the 
observation  that  the  absence  of  remains  of  these  ground-sloths 
from  the  Miocene  of  North  America,  coupled  with  their  presence 
in  the  Santa  Cruz  beds  of  Patagonia,  clearly  indicates  that  they  are 
late  immigrants  from  the  south  into  the  northern  half  of  the 
continent.  , 

Nearly  allied  to  Mylodon,  the  genus  Glossotherium  from  the 
Plistocene  of  Argentina  and  Uruguay  serves  to  connect  it  with 
another  generic  representative  of  the  family  known  as  Scelido- 
therium. In  place  of  the  comparatively  short  skulls  of  the 
mylodons,  the  species  of  this  genus  have  the  muzzle  of  the  skull 
greatly  elongated,  so  that  there  is  a  long  toothless  space  in 
advance  of  the  dental  series ;  and  whereas  the  skulls  of  the  species 
of  Mylodon  are  essentially  sloth-like,  those  of  Scelidotherium  show 
a  marked  approximation  to  the  anteater-type.  The  species  of 
Scelidotherium  are  of  medium  or  rather  small  size ;  and  in  space 
the  genus  ranges  from  Patagonia,  through  the  Argentine,  to  Brazil, 
Bolivia,  and  Chili ;  while  in  time  it  extends  from  the  superficial 
sand-dunes  and  Pampean  deposits  to  the  lower  Tertiaries  of 
Parana,  Monte  Hermoso,  Catamarca,  and  Santa  Cruz,  with  a 
gradual  decrease  in  the  size  of  the  species  as  we  descend  in  the 
geological  scale '.  Nearly  allied  is  the  genus  Catonyx,  from  the 

1  The  Santa  Cruz  form  has  been  quite  unnecessarily  separated  under  the 
name  of  Analcithermm. 


io6 


THE   NEOG^EIC   REALM. 


[CHAP. 


! 


w     ^ 

K^*  •-£ 


III.]  MARSUPIALS.  107 

Brazilian  caverns ;  while  Nothrotherium  of  the  same  deposits 
seems  to  have  been  another  nearly  related  form  with  teeth  of  the 
Megalotherium  type.  The  imperfectly  known  Nothropus  of  the 
Pampean  and  Ortotherium  of  the  Parana  beds  seem,  on  the  other 
hand,  to  be  late  survivals  of  another  group  typically  represented 
by  the  genera  Eucholaops  and  Pseudhapalops  of  the  Santa  Crucian 
epoch  of  Patagonia.  These  forms  differ  from  all  those  noticed 
above  in  that  the  terminal  joints  of  some  of  the  toes  have  a 
median  cleft  as  in  the  great  anteater,  and  likewise  in  the 
elongation  of  the  metatarsal  bones  ;  and  it  seems  probable  that  the 
hind  foot  was  not  so  much  everted  as  in  the  later  representatives 
of  the  family.  The  skull  is  of  the  general  type  of  that  of  Mylodon ; 
most  of  the  molars  being  prismatic  in  form,  and  surmounted  by  a 
pair  of  transverse  ridges,  more  or  less  closely  connected  at  their 
extremities  so  as  to  produce  an  oval  cavity  on  the  grinding  surface. 
The  first  tooth  is,  however,  tusk-like,  and  separated  by  a  gap  from 
the  others.  In  some  of  these  early  ground-sloths  the  skull  did 
not  exceed  three  inches  in  length ;  but  other  species  were  con- 
siderably larger.  They  are  evidently  generalised  types,  and  were 
probably  nearly  allied  to  the  ancestral  stock  which  gave  rise  to 
Mylodon  and  Megalonyx,  if  indeed  they  be  not  the  actual  pro- 
genitors of  both. 

The  last  group  for  consideration  is  that  of  the  marsupials,  or 
pouched  mammals,  among  which  the  family  of  the 
opossums  (Didelphyidce),  with  the  three  genera 
Didelphys1,  Dromiciops,  and  Chironectes,  is  now  confined  to  the 
New  World,  the  great  majority  of  the  numerous  species  being 
Neogaeic,  although  the  common  opossum  (Didclphys  marsupialis} 
ranges  from  Chili  and  Brazil  to  the  United  States.  Although 
certain  forms  from  the  Santa  Cruz  beds  described  under  the 
names  of  Eodidelphys  and  Prodidclphys  were  originally  assigned  to 
the  present  family,  these  have  been  subsequently  identified  t^y 
Dr  Ameghino2  with  the  under-mentioned  family  of  the  Microbio- 
theriidce.  True  opossums  occur,  however,  in  the  Monte  Hermoso 

1  This  genus  is  divided  into  several  sub-genera,  regarded  by  some  writers  as 
entitled  to  generic  rank. 

2  Bol.  Ac.  Cordoba,  Vol.  xm.  p.  363  (1894). 


io8 


THE    NEOG^IC   REALM. 


[CHAP. 


beds  •  while,  as  mentioned  in  the  last  chapter,  they  were  widely 
distributed  in  the  northern  hemisphere  during  the  Oligocene.  If 
the  conclusions  of  Dr  Ameghino  are  right  as  to  the  absence  of 
these  marsupials  from  the  Santa  Cruz  beds,  it  is  evident  that 
opossums  only  reached  South  America  from  the  north  at  the  close 
of  the  Miocene  or  commencement  of  the  Pliocene,  and  that  they 
do  not  belong  to  the  indigenous  fauna.  It  has  been  generally 
considered  that  the  common  opossum  of  the  United  States  is  a 
direct  survivor  from  the  Oligocene  forms  of  North  America,  but  it 
is  more  probable  that  it  is  really  a  very  recent  immigrant  from  the 
south,  seeing  that  fossil  representatives  of  the  genus  are  unknown 
from  the  North  American  Miocene  and  Pliocene.  During  the 
Miocene  the  group  perhaps  survived  in  the  extreme  south  of 
North  America. 

Although  opossums  are  apparently  wanting,  the  Santa  Cruz 
beds  have  yielded  remains  of  undoubted  marsupials,  but  several 


FIG.  25.     LEFT  HALF  OF  LOWER  JAW  OF  Prothylctcinus, 


nat.  size.) 


of  them  are  assigned  by  Dr  Ameghino  to  a  distinct  group  under 
the  name  of  Sparassodonta.  Foremost  of  these  is  the  genus 
ProthylacinuS)  already  mentioned  in  the  last  chapter,  which  may 
be  provisionally  assigned  to  the  Australian  Dasyuridce.  In  having 
only  three  in  place  of  four  lower  pairs  of  incisors  this  genus  agrees 
with  the  latter  family,  and  differs  from  the  opossums  ;  while  the 
whole  character  of  the  lower  jaw  and  dentition  is  very  similar  to 
that  of  the  Tasmanian  Thylatinus,  with  the  exception  that  the 
premolars  are  closer  together.  As  in  the  Dasyuridce  generally, 


III.]  MARSUPIALS.  109 

there  are  four  pairs  of  upper  and  three  of  lower  incisor  teeth  in 
Prothylacinus,  and  the  same  is  the  case  with  the  smaller  Santa 
Crucian  form  described  as  Amphiproviverra,  which  appears  to  be 
of  a  distinctly  dasyurid  type,  although  not  coming  very  near  to 
any  Australian  genus.  v 

With  regard  to  the  Microbiotheriida,  as  typified  by  the  genus 
Microbiotherium,  these,  although  they  are  not  included  by  Dr 
Ameghino  in  the  order,  appear  to  be  undoubted  marsupials,  since 
they  have  a  dentition  numerically  the  same  as  that  of  the  opossums, 
vacuities  in  the  palate,  and  an  inflected  angle  to  the  lower  jaw. 
From  the  opossums  they  differ  by  the  non-production  of  the  palate 
behind  the  last  molars,  and  in  the  form  of  the  lower  jaw,  in  which 
the  extremity  is  produced  to  a  greater  extent  in  advance  of  the 
canine.  In  all  these  respects  they  approximate  to  the  Dasyurid 
genus  Phascologale,  from  which  they  differ  in  having  one  pair  less 
of  incisors  in  each  jaw.  The  ancestors  of  the  Australian  Dasyuridce. 
must,  however,  have  originally  had  five  pairs  of  upper  and  four  of 
lower  incisor  teeth,  as  the  former  are  retained  in  many  of  the  ban- 
dicoots (Peramelida),  while  Myrmecobms  occasionally  develops  four 
pairs  of  these  teeth  in  the  lower  jaw.  It  seems  therefore  probable 
that  the  Microbiotheriidcz  were  minute  polyprotodont  marsupials 
of  an  Australian  type. 

There  is  more  difficulty  in  arriving  at  any  satisfactory  con- 
clusions as  to  the  serial  position  of  certain  carnivorous  mammals 
from  the  Santa  Cruz  beds,  of  which  a  large  form  described  as 
Borhycena  may  be  taken  as  an  example.  In  these  animals  the 
dentition  approximates  to  a  certain  extent  to  that  of  the  primitive 
or  creodont  Carnivora  of  the  earlier  Tertiaries  of  the  northern 
hemisphere,  although  retaining  the  marsupial  feature  of  four  pairs 
of  molars  and  only  three  of  premolars.  The  replacement  of  the 
teeth  is  also  fuller  than  in  the  marsupials.  Dr  Ameghino  has 
suggested  that  these  animals  were  transitional  between  marsupial 
and  eutherian  carnivores,  and  that  the  latter  group  originated  in 
South  America ;  but  this  idea  is  obviously  untenable.  A  possible 
suggestion  is  that  they  may  be  specialised  offshoots  from  the 
marsupial  stock  which  died  out  without  giving  origin  to  any 
descendants. 

A  small  mouse-like  mammal  first  described  in  1863  upon  the 


110  THE    NEOG^IC  REALM.  [CHAP. 

evidence  of  a  single  specimen  from  Ecuador  under  the  name  of 
Hyracodon  fuliginosus,  but  whose  affinities  were  not  determined 
till  1895,  when  an  example  of  a  second  and  larger  species  was 
procured  from  Bogota,  belongs  to  the  sole  surviving  genus  of  a 
group  of  small  marsupials  which  occur  abundantly  in  the  Santa 
Cruz  beds,  and  were  till  quite  recently  regarded  as  extinct.  Un- 
fortunately the  name  Hyracodon  has  been  previously  employed 
to  designate  an  extinct  ungulate,  and  it  has  accordingly  been 
replaced  by  Ccenolestes.  The  essential  characteristic  of  this  group 
of  marsupials,  is  that  while  their  upper  dentition  is  of  a  poly- 
protodont  type,  that  of  the  lower  jaw  is  very  similar  to  the 
diprotodont  modification.  In  the  living  species,  for  instance, 
there  are  four  pairs  of  small  upper  incisors  of  a  normal  type, 


FlG.    26.      FORE    PART    OF   THE    RIGHT    HALF    OF   THE   LOWER  JAW    OF 

Acdestis  oweni.     (Much  enlarged.) 

The  first  tooth  on  the  right  is  the  first  incisor,  and  the  one  on  the  extreme 
left  the  first  molar. 

followed  by  a  large  canine,  while  in  the  lower  jaw,  as  shown  in 
the  accompanying  figure  of  that  of  one  of  the  extinct  forms,  there 
is  a  large  pair  of  horizontally  projecting  incisors,  succeeded  by 
several  minute  functionless  teeth,  of  which  the  first  three  represent 
the  second  and  third  incisors  and  the  canine.  In  all  the  forms 
the  molar  teeth  have  quadrangular  crowns,  surmounted  by  four 
blunt  tubercles,  somewhat  resembling  in  structure  those  of  certain 
ungulates,  and  thus  totally  different  from  the  triangular  and 
sharply-cusped  molars  of  the  opossums  and  other  polyprotodonts. 
In  the  living  forms,  as  well  as  in  certain  fossil  kinds  (Epanorthus, 
Decastis  and  Acdestis]  from  the  Santa  Cruz  beds,  the  last  premolar 
tooth,  as  shown  in  the  figure  of  the  jaw  of  Acdestis,  is  of  normal 
dimensions  :  and  these  forms  may  consequently  be  grouped  in  a 
single  family,  under  the  name  of  Epanorthidce.  In  another  group, 


III.]  MARSUPIALS.  1 1 1 

confined  to  the  Santa  Crucian  horizon,  where  it  is  represented  by 
the  family  Abderitidce,  the  last  premolar  in  each  jaw  is  much 
larger  and  taller  than  the  other  teeth,  and  has  its  crown  in  the 
form  of  a  compressed  cone,  marked  on  the  sides  with  vertical 
grooves,  as  exhibited  in  the  figure  of  Abderites.  A  third  family  is 


FlG.    27.       RIGHT    HALF   OF    LOWER  JAW   OF   Abderites,    MUCH    ENLARGED. 

known  as  the  Garzoniidce.  In  all  the  skull  is  of  an  elongated 
form,  with  large  vacuities  both  in  the  front  and  hinder  part  of  the 
palate,  and  presents  a  considerable  general  resemblance  to  those 
of  the  Australian  genera  Peragale  and  Perameles.  With  the  ex- 
ception of  the  retention  of  four  pairs  of  upper  incisors  and  the  small 
size  of  all  these  teeth,  the  dentition  exhibits,  however,  a  remark- 
able approximation  to  that  of  the  Australian  diprotodont  genus 
Dromicia.  On  the  other  hand,  the  feet  are  of  normal  structure, 
with  five  complete  toes,  none  of  which  are  united  by  integuments; 
the  thumb  and  great  toe  being  apparently  slightly  opposable  to 
the  other  digits.  Probably  the  rat-like  tail  is  slightly  prehensile 
at  the  extremity ;  and  a  small  pouch  is  present  in  the  female.  In 
the  skeleton  the  lower  jaw  exhibits  the  usual  inflection  of  the 
angle ;  and  the  pelvis  carries  marsupial  bones. 

Probably  these  Patagonian  marsupials,  which  may  be  known 
as  selvas,  must  be  included  in  the  diprotodont  sub-order ;  from 
the  Australian  representatives  of  which  they  differ  by  the  small 
and  numerous  upper  incisors  and  the  non-syndactylous  hind  feet. 
Both  these  being  generalised  features,  it  is  evident  that  if  the 
selvas  are  true  diprotodonts  their  ancestors  must  have  originated 
from  the  polyprotodonts  in  Notogaea,  for  if  they  are  of  exclusively 
South  American  origin  they  must  form  a  subordinal  group  by 


112  THE    NEOG^IC   REALM.  [CHAP. 

themselves.  Assuming  their  affinities  with  the  Australian  type 
to  be  rightly  determined,  they  constitute  a  most  important  link 
in  the  chain  connecting  the  faunas  of  South  America  and  Aus- 
tralia. 

In  the  last  chapter  it  has  been  argued  that,  from  the  absence 
of  allied  forms  in  the  Tertiaries  of  North  America  and  Europe,  as 
well  as  from  their  resemblance  to  the  Australian  dasyurids,  it  is 
difficult  to  come  to  any  conclusion  other  than  that  the  ancestors  of 
the  Santa  Crucian  polyprotodont  marsupials  reached  the  country 
from  Australia,  either  by  way  of  the  Antarctic  continent,  or  by  a 
land-bridge  in  a  more  northern  part  of  the  Pacific.  If  this  be 
correct,  and  likewise  the  supposition  that  the  opossums  origi- 
nated from  the  ancestral  stock  in  South-eastern  Asia,  it  will  be 
evident  that  Didelphys  and  Ccenolestes  met  in  South  America 
after  their  ancestors  had  travelled  half  round  the  world  in  opposite 
hemispheres. 

It  may  be  added  that  the  alleged  occurrence  of  monotremes 
in  the  Santa  Cruz  beds  is  due  to  bones  of  aberrant  armadillos 
(Peltephilus)  having  been  mistaken  for  those  of  that  group1. 

Although  in  this  volume  the  writer  avoids  laying  much  stress 
upon  aquatic  mammals,  it  may  be  mentioned  that 

Cetaceans. 

there  are  two  genera  ot  dolphins  belonging  to  the 
family  Platanistidce,  each  represented  by  a  single  species,  which  are 
peculiar  to  the  Neogaeic  realm.  These  are  Stenodelphis  (Pontoporia] 
from  the  mouth  of  the  Rio  de  la  Plata,  and  Inia  of  the  upper 
Amazon;  the  only  other  existing  representative  of  the  family 
being  Platanista  of  the  larger  Indian  rivers. 

After  the  foregoing  survey  of  the  chief  features  of  the  recent 

and  fossil  mammalian  fauna  of  the  Neogaeic  realm, 
tinction  ofThe  ^ts  general  bearings  on  the  relations  of  South  America 

to  other  parts  of  the  world  may  be  taken  into  con- 


sideration. It  will,  however,  facilitate  matters  to 
give  a  tabular  view  of  the  orders,  suborders,  and  families  of  non- 
volant  land  mammals  represented  in  the  realm.  In  the  following 
table  such  groups  as  are  either  confined  to  Neogsea,  or  have  only 
reached  North  America  at  a  comparatively  recent  epoch  are 

1  See  Lydekker,  An.  Mus.  La  Plata—  Pal.  Argent.  Pt.  III.  p.  67  (1894). 


III.]  LIST   OF   THE   FAUNA.  113 

printed  in  italics ;  the  extinct  types  being  indicated  by  the  prefix 
of  an  *,  while  those  dating  from  the  Santa  Crucian  (or  the  earlier 
Patagonian)  epoch  are  followed  by  the  words  Santa  Cruz,  and 
those  from  the  Parana  beds  by  the  word  Parana. 

I.     PRIMATES. — Santa  Cruz. 

Cebida, — Santa  Cruz  (* Hdmunculus). 
Hapalidcz. 

II.     CHIROPTERA. 

Phyllostomatida. — One    genus  ranging   to    Cali- 
fornia. 

Emballonuridae. — Seven  peculiar  genera. 
Vespertilionidae. — Natalus,   Thyroptera. 

III.  INSECTIVORA. 

Solenodontidce. — West  Indies. 

IV.  CARNIVORA. 

Felidae. — No  peculiar  genera. 

Canidae. — In  addition  to  the  cosmopolitan  Canis, 

represented     in     Brazil     by    the     peculiar 

Icticyon. 
Ursidae. — No    peculiar   genera;    *Arctotherium, 

common  to  N.  America. 
Procyonidae — Nasua,  Cercoleptes,  Bassaricyon,  and 

*  Cynonasua. 
Mustelidae. — Galictis,   and    Conepatus,  the  latter 

extending  into  Texas. 

V.     UNGULATA. 

i.     ARTIODACTYLA. 

Cervidae. — Cariacus,  peculiar  to  New  World. 
Dicotylidae. — Peculiar  to  New  World  at  present 

day. 

Camelidae. — Lama. 
L.  8 


114  THE    NEOG^IC   REALM.  [CHAP. 

2.  PERISSODACTYLA. 

Tapiridae. — Elsewhere  only  in  Malaysia  at  the 
present  day,  but  formerly  widely  distributed 
over  the  northern  hemisphere. 

Equidae. — Now  unknown,  except  through  intro- 
duction. In  addition  to  the  cosmopolitan 
Equus  (including  Tertiary  forms),  the  pecu- 
liar Pampean  genera  *  Hippidium  and  *  Ono- 
hippidium. 

3.  *  Litopterna.— Santa  Cruz. 

* Macraucheniidce. — Santa  Cruz. 

*  Proter other iidtz. — Santa  Cruz. 

4.  *  Astrapotheria. — Santa  Cruz. 

*  A strapotheriidce. — Santa  Cruz. 

* Homalodontotheriida. — Santa  Cruz. 

5.  *  Toxodontia. — Santa  Cruz. 

*  Toxodontidce. — Santa  Cruz. 

*  Typotheriida. — Santa  Cruz. 

*  Pachyruchidce. — Santa  Cruz. 

6.  * Pyrotheria. — Patagonian  beds  lying  below  those  of 

Santa  Cruz. 

*  Pyrotheriidte. — Patagonian  beds. 

7.  PROBOSCIDEA. 

Elephantidse. — Represented  by  Mastodon  in 
the  Pampean  and  Monte  Hermoso  beds. 

VI.       RODENTIA. 

I.       SCIUROMORPHA. 

Sciuridae. — Represented  by  Sciurus  as  far  south 
as  Paraguay. 

*  Castoroididse. — Peculiar  to  New  World. 

2      MYOMORPHA. 

Muridae. — Represented  by  species  of  the  New 
World  genus  Sitomys,  as  well  as  by  several 
peculiar  types  such  as  Rhithrodon,  Ichthyomys 
of  Peru,  Holochilus  of  Brazil,  etc. 


III.]  LIST   OF   THE   FAUNA.  115 

3.  HYSTRICOMORPHA. — Santa  Cruz. 

Caviidce. — Paran  a. 
Dasyproctidce . 
Dinomyidce. 

Lagostomatidcz. — Santa  Cruz. 
Hystricidae. — Santa  Cruz. 

Octodontidae. — Mainly  Neotropical,  but  also  Ethi- 
opian.    Santa  Cruz. 

*  Eocardiidce. — Santa  Cruz. 

4.  LAGOMORPHA. 

Leporidae. — Represented     by    two     species     of 
Lepus. 

VII.      EDENTATA. — Santa  Cruz. 

Dusypodida. — Santa  Cruz.  A  few  forms  range 
as  far  as  Texas. 

*  Glyptodontida. — Santa  Cruz.    One  Neogaeic  genus 

ranges  as  far  north  as  Texas,  and  a  peculiar 
one  has  been  described  from  further  north. 

*  Megalotheriidce. — Santa  Cruz.     Mainly  Neogaeic, 

but  also  ranging  into  North  America. 
Myrmecophagidcz. 
Bradypodidcz. 

VIII.     MARSUPIALIA. — Santa  Cruz. 

1.  DIPROTODONTIA. — Santa  Cruz.     (Elsewhere  only  in 

Notogaea). 

Epanorthidcz.  Santa  Cruz.  (One  existing  genus, 
Ccenolestes). 

*  Abderitidcz. — Santa  Cruz. 

*  GarzoniidcB. — Santa  Cruz. 

2.  POLYPROTODONTIA. — Santa  Cruz. 

Didelphyidae. — Now  mainly  Neogaeic,  where  they 
date  from  the  Monte  Hermoso  stage,  but 
ranging  into  North  America,  and  formerly 
widely  spread  over  the  northern  hemisphere. 
Chironectes,  several  subgenera  of  Didelphys, 
and  Dromiciops  peculiar  to  the  realm. 

8—2 


Il6  THE    NEOG^EIC    REALM.  [CHAP. 

VIII.     MARSUPIALIA  (continued}. 

Dasyuridae. — Santa  Cruz.  Now  confined  to 
Notogaea,  but  apparently  represented  in  the 
Santa  Cruz  beds  by  * Prothylacinus  and 
*  Amphiproviverra. 

*  Microbiotheriidcz. — Santa  Cruz. 
Incertce,  Seats. 

*  Borhyanidce. — Santa  Cruz. 

Although  the  addition  of  the  names  of  all  the  peculiar  genera, 
both  recent  and  extinct,  would  have  rendered  the  distinctness  of 
the  Neogaeic  mammalian  fauna  still  more  pronounced,  yet  the 
foregoing  table  as  it  stands  is  amply  sufficient  to  show  that 
Neogaea  is  entitled  to  form  one  of  the  three  primary  zoological 
realms  of  the  world.  Starting  with  the  Santa  Crucian  epoch  of 
Patagonia  and  the  somewhat  older  Patagonian  stage,  which  form 
the  earliest  date  at  which  the  history  of  the  Tertiary  land  mammals 
of  Neogaea  can  be  taken  up,  there  is  evidence  that  at  least  the 
southern  part  of  the  area  was  populated  by  the  following  pecu- 
liar fauna.  Firstly,  monkeys  of  a  type  quite  different  from  those 
of  the  Old  World,  but  evidently  allied  to  the  existing  Neogaeic 
forms,  were  abundant;  while  rodents,  belonging  to  the  same 
groups  as  those  now  inhabiting  the  continent,  several  of  which 
were  nearly  allied  to  existing  African  forms,  and  more  remotely  to 
certain  Oligocene  European  types,  attained  a  great  development. 
Probably  Insectivora  with  V-shaped  molars  were  also  present. 
More  peculiar  are  the  extinct  subordinal  groups  of  ungulates 
described  above,  which  appear  to  have  been  allied  to  the  ancestors 
of  the  perissodactyles  of  the  northern  hemisphere,  and  may 
possibly  be  remotely  connected  with  the  African  hyraces.  At  the 
same  period  flourished  several  families  of  edentates  (a  group 
which  in  its  restricted  sense  was  originally  peculiar  to  Neogaea), 
such  as  armadillos,  glyptodonts,  and  ground-sloths,  most  of  the 
members  of  which  were  of  comparatively  small  size ;  but  of  the 
ancestry  of  this  group  nothing  can  be  said  with  certainty. 
Among  the  marsupials,  although  opossums  appear  to  have  been 
wanting,  there  were  several  types  seemingly  allied  to  Notogaeic 
forms,  while  others  which  may  be  included  in  the  order  were  more 


III.] 


RELATIONS   OF   THE   FAUNA. 


117 


or  less  unlike  any  from  other  regions.  In  addition  to  true  mar- 
supials, the  only  carnivorous  types  were  the  problematical 
Borhycenidce. 

Now,  as  stated  in  the  earlier  part  of  the  chapter  (p.  68), 
this  fauna  cannot  be  older  than  the  lowest  Miocene  or  highest 
Oligocene ;  and  among  its  deficiencies  may  be  noted  lemuroids,  true 
carnivores,  creodonts,  artiodactyle  and  perissodactyle  ungulates, 
and  opossums,  all  of  which  were  in  existence  during  the  Oligo- 
cene or  Miocene  in  North  America  and  Europe.  Moreover,  at  those 
epochs  the  former  country  lacked  the  whole  of  the  Neogaeic  types. 

Clearly,  then,  there  must  have  been  a  barrier  between  North 
and  South  America  during  the  Oligocene  and  a 

Early  Separ- 

portion  or  the  whole  of  the  Miocene;  but  before  ationofN.and 
entering  into  the  consideration  of  other  evidence  S'  Amenca- 
showing  the  nature  of  that  barrier,  it  may  be  well  to  give  a  table  of 
the  mammaliferous  Tertiary  strata  of  North  and  South  America, 
with  their  approximate  European  equivalents1.  In  descending 
order  this  runs  as  follows  : 


Plistocene 
Up.  Pliocene 
Low.  Pliocene 

Miocene 

Up.  Oligocene 

Mid.  Oligocene 


Low.  Oligocene 
Up.  Eocene 
Mid.  Eocene 

Low.  Eocene 
Lowest  Eocene 


South  America. 
Pampean 


Santa  Cruz 
Patagonian 


North  America. 
Equus  beds. 
Blanco 

Monte  Hermoso  f  Palo  Duro. 

(?)  Parana  Loup-Fork  -j  Nebraska 

tDeep  River 

(Hiatus) 
John  Day 

/  Protoceras 
Beds 

White  River  j°reodon 
Beds 


Titanother- 


ium  Beds 
Uinta 

rWashakii 
Bridger  -I  Bridger. 

IWind  River. 
Wahsatch 
Puerco 


Europe. 

Cave-deposits  etc. 
?  Crag. 
|  Pikermi 

Sansan 

St  Gerand-Le-Puy 


Ronzon. 

Montmartre 
Parisian 

Suessonian 
Cernaysian 


1  In  compiling  this   table  the  writer  is  indebted  to  Prof.   W.   B.   Scott. 
Many  American  geologists  (among  them  Dr  Scott)  include  the  whole  of  the 


Il8  THE    NEOG^IC   REALM.  [CHAP. 

Regarding  the  geological  evidence  of  a  separation  between  the 
two  Americas,  Cretaceous  marine  strata  occupy  a  large  portion  of 
Mexico;  and  in  1879  Dr  Le  Conte1  wrote  as  follows.  The  shore 
line  of  the  Gulf  of  Mexico  "was  much  more  extended  both  north- 
ward and  westward  than  either  now  or  in  Tertiary  times.  From 
the  Gulf  there  extended  northwestward  an  immensely  wide  sea, 
covering  the  Plains  region  and  the  Rocky  Mountain  region  as  far 
westward  as  the  Wahsatch  range,  and  dividing  the  continent  into 
two  continents,  an  eastern  or  Appalachian,  and  a  western  or 
Basin-region  continent.  Probably  also  this  sea  connected  across 
the  region  of  Mexico  with  the  Pacific,  thus  dividing  the  western 
continent  into  two,  a  northern  and  southern."  Later  observations 
have  shown  that  the  Cretaceous  sea  undoubtedly  made  a  wide  gap 
between  North  America  and  the  southern  portion  of  the  con- 
tinent2; while  the  existence  of  Oligocene  or  Miocene  strata  in  the 
region  of  the  isthmus  of  Parana  shows  that  the  separation,  which 
probably  continued  through  the  Eocene,  was  in  existence  during 


Loup-Fork  in  the  Miocene;  while  to  the  lower  part  of  the  same  era  they 
assign  the  John  Day  beds  of  America,  and  the  St  Gerand-le-Puy  beds  of 
Europe.  Others  (e.g.  Prof.  Osborn,  Studies  Biol.  Labor.  Columbia  Coll. 
vol.  i.  pt.  2,  p.  28,  1893)  refer  the  Equus  beds  of  North  America  to  the 
Pliocene.  The  following  quotation  from  a  paper  by  Prof.  Cope  (American 
Naturalist,  1895,  p.  599)  conclusively  proves  their  Plistocene  age.  There  it 
is  stated  that  "the  Equus  beds  are  found  covering  areas  of  various  extent  in 
Oregon,  Nevada,  California,  the  Staked  Plains,  Southern  Texas,  Chihuahua 
and  the  valley  of  Mexico.  Their  most  eastern  station  is  western  Nebraska. 
They  contain  a  fauna  which  includes  one  extinct  species  (Equus  major  Dek.)  of 
the  Megalonyx  fauna,  and  the  recent  Castor  fiber.  They  contain  the  extinct 
genus  of  sloths  Mylodon,  of  a  species  different  from  that  of  the  east,  and  four 
species  of  camels  of  the  extinct  genus  ffolomeniscns,  and  a  peccary.  Recent 
species  of  Cam's  and  Thomomys  occur,  while  two  extinct  horses  (Equus  occi- 
dentalis  Leidy  and  E.  tau  Owen)  are  common.  The  hairy  elephant  (E.  primi- 
genius)  is  abundant,  while  the  Mastodon  americanus  is  rare,  if  occurring  at  all. 
The  proportion  of  recent  to  extinct  species  and  genera  in  the  Equus  bed  fauna 
is  very  similar  to  that  occurring  in  the  Megalonyx  fauna,  while  they  differ  as  to 
details." 

1  Elements  of  Geology,  pp.  451,  452,  New  York  (1879). 

2  This    separation   also  existed  in  the  Jurassic  era,   when,  as  shown  by 
Neumayr  (Erdgeschichte,  2nd  ed.  vol.  II.,  p.   263),  South  America  was  united 
across  the  Atlantic  area  with  Africa  and  Madagascar. 


I 

III.]  RELATIONS   OF   THE   FAUNA.  119 

the  middle  portion  of  the  Tertiary  epoch1.  When  the  connection 
between  North  and  South  America  was  completed  is  not  precisely 
fixed  by  geological  evidence  ;  but  the  occurrence  of  a  glyptodont 
in  the  Nebraska  stage  of  the  Loup-Fork  group,  shows  that  it  must 
have  been  by  the  end  of  the  Miocene2.  The  question  of  a  con- 
nection between  the  two  continents  by  way  of  the  West  Indies 
is  discussed  later  in  this  chapter,  where  it  is  concluded  that  if 
such  a  connection  existed  at  all,  it  must  have  been  of  a  transient 
nature. 

Having  thus  shown,  both  from  palaeontological  and  geological 
evidence,  that  the  early  mammalian  fauna  of 

Incursion  of 

Neogaea  appears  to  have  been  totally  isolated  from  Northern 
that  of  North  America  up  to  about  the  end  of  the 
Miocene,  the  question  of  the  origin  of  that  fauna  may  be  deferred, 
and  the  irruption  of  northern  types  after  the  connection  between 
North  and  South  America  had  been  established  taken  into  con- 
sideration. It  may  be  mentioned,  however,  that  it  was  not  till 
after  this  irruption  of  the  northern  forms  that  the  essentially 
Neogaeic  fauna  attained  its  maximum  development  in  respect  to 
the  bodily  size  of  its  constituents ;  since  a  gradual  increase  in  this 
respect  maybe  traced  from  the  small  glyptodonts  and  sloths  of  the 
Santa  Cruz  epoch,  through  the  larger  ones  of  the  Monte  Hermoso 
horizon,  to  the  gigantic  forms  characteristic  of  the  Pampean  and 
the  cavern  deposits  of  Brazil. 

The  presence  of  a  glyptodont  in  the  Nebraska  stage  of  the 
Loup-Fork  group  in  North  America,  and  of  northern  forms  in  the 
Monte  Hermoso  horizon  of  South  America,  marks,  then,  the  first 
commingling  of  the  original  faunas  of  the  two  halves  of  the  New 
World.  For  the  first  time  in  the  history  of  the  southern  continent 
this  connection  allowed  of  the  immigration  from  the  north  of  the 
true  Carnivora,  such  as  the  existing  cats  (Felts)  the  extinct  sabre- 
toothed  tigers  (Macharodus),  dogs  and  foxes  (Canida),  bears 
(Ursus  and  Arctotherium\  raccoons  (Procyonidtz),  skunks  and 
their  allies  (Mustetida),  together  with  various  ungulates  belonging 
to  sub-orders  previously  unknown  in  the  realm.  These  latter 
include  the  guanaco  and  vicuna  (Lama] — of  which  ancestral  forms 

1  See  J.  W.  Gregory,  Quart.  Joum.  GeoL  Soc.  Vol.  LI.  pp.  299,  300  (1895). 

2  As  stated  above,  many  refer  the  whole  Loup-Fork  group  to  the  Miocene. 


I2O  THE    NEOG^EIC   REALM.  [CHAP. 

are  abundant  in  the  North  American  Tertiaries — New  World  deer 
(Cariacus],  horses  (Equidce)  of  various  genera,  tapirs  (Tapirid&\ 
peccaries  (Dicotylidte),  and  mastodons.  Among  the  rodents, 
squirrels,  the  various  genera  of  Muridce,  and  the  hares,  likewise  at 
this  epoch  made  their  first,  appearance  on  the  scene.  Opossums 
also  at  this  time  effected  an  entrance  into  the  land  which  has  now 
become  their  chief  home.  That  this  new  fauna  came  in  from 
North  America,  and  not  from  any  other  part  of  the  world,  may  be 
regarded  as  certain  from  the  presence  of  such  essentially  New 
World  types  as  raccoons  and  their  allies,  skunks,  peccaries, 
Cariacus,  and  Camelidce  (exclusive  of  the  Old  World  genus 
Came/us,  which  is  of  late  origin),  coupled  with  the  absence  of 
true  deer  (Cervus),  pigs  (Sus),  Old  World  monkeys,  and  lemurs. 

At  the  same  time  this  union  of  the  northern  and  southern 
halves  of  the  New  World  allowed  certain  members  of  the  original 
Neogaeic  fauna  to  make  their  escape  into  North  America,  glypto- 
donts,  as  already  said,  making  their  appearance  in  the  Nebraska 
stage  of  the  Loup-Fork  group  of  the  United  States,  while  the 
ground-sloth  Megalonyx  occurs  in  the  Blanco  Beds. 

Although  it  is  not  universally  admitted1,  there  is  some 
evidence  to  indicate  that  this  land  connection  was  of  com- 
paratively brief  duration,  seeing  that  none  of  the  characteristic 
extinct  types  of  South  American  ungulates,  nor  any  of  the  peculiar 
Neogaeic  rodents,  reached  the  northern  half  of  the  continent. 

During  the  whole  time  that  the  alluvial  deposits  of  the  Parana 
and  Paraguay  rivers  were  being  laid  down,  and  well  on  into  the 
human  period,  the  mammalian  fauna  of  the  Pampean  epoch,  formed 
by  an  admixture  of  southern  and  northern  types,  continued  to 
flourish,  until  the  time  when  there  came  a  complete  sweep  of  all 
the  larger  forms,  clearing  off  the  whole  of  the  ground-sloths, 
glyptodonts,  mastodons,  toxodonts,  macrauchenias,  horses,  sabre- 
toothed  tigers,  and  the  larger  members  of  the  camel  tribe,  and  in 
the  Argentine  leaving  only  armadillos,  guanacos,  a  few  deer,  a 
number  of  rodents,  various  cats  and  foxes,  as  well  as  skunks  and 
certain  other  members  of  the  weasel  family,  to  represent  the  vast 
assemblage  of  strange  and  giant  creatures  that  once  roamed  over 

1  See  Gregory,  op.  cit.  p.  300. 


III.]  DISAPPEARANCE   OF   LARGE   FORMS.  121 

its  plains.  With  regard  to  this  extraordinary,  and  apparently 
sudden  disappearance  of  almost  all  the  larger  forms  of  animal  life 
from  South  America,  it  may  be  pretty  confidently  asserted,  from 
the  organisation  of  the  creatures  themselves,  that  at  the  time  when 
the  ground-sloths  flourished,  extensive  portions  of  what  is  now  the 
open  pampas  of  Argentina  were  covered  with  forest ;  and  why  the 
whole  district  should  have  become  practically  treeless,  seeing  that 
trees  like  the  Australian  eucalypti  grow,  when  introduced,  with 
more  vigour  than  in  their  native  home,  is  exceedingly  hard  to 
understand.  That  the  country  even  when  thus  denuded  was 
unsuited  to  the  needs  of  the  larger  forms  of  mammalian  life,  is, 
however,  negatived  by  the  circumstance  that  in  many  parts  the 
horses  and  cattle  introduced  from  Europe  have  run  wild  and 
increased  to  an  almost  unprecedented  extent.  Neither  does  it 
appear  that  the  extermination  can  be  attributed  to  a  period  of 
extreme  cold,  since  a  glaciation  of  the  pampas  would  assuredly 
have  left  unmistakable  evidence  of  its  presence.  It  is  likewise 
practically  certain  that  the  clean  sweep  of  the  forests  of  Argentina 
and  the  larger  mammals  of  the  whole  of  South  America  is  not  due 
to  the  hand  of  man.  It  has,  indeed,  been  suggested  that  the  vast 
herds  of  guanaco  which  formerly  roamed  the  pampas  may  have 
cleared  the  forests  by  preventing  the  growth  of  the  seedlings ;  but 
when  we  recall  the  fact  that  numbers  of  this  group  of  animals 
flourished  during  the  period  when  the  alluvial  formation  was  in  the 
course  of  being  deposited,  it  scarcely  looks  as  though  this  could 
have  been  a  vera  causa.  Moreover,  if  the  forests  were  by  some 
means  or  other  actually  destroyed,  and  the  extermination  of  their 
animal  denizens  thus  encompassed,  there  would  still  remain  the 
disappearance  of  plain-dwelling  forms,  like  horses,  to  be  accounted 
for.  Some  have  thought  that  pumas,  by  preying  on  the  colts, 
were  the  active  agents  in  this  instance  ;  but  even  if  such  were 
really  the  case,  it  gives  no  help  with  regard  to  the  disappearance 
of  the  ground-sloths  and  glyptodonts; — the  latter  being  such 
strongly  armoured  creatures  that  it  is  absolutely  certain  they  were 
not  killed  off  by  any  animal  foes.  The  problem  is  further  com- 
plicated by  the  circumstance  that  the  fossil  remains  of  nearly  all 
the  larger  animals  which  formerly  inhabited  the  pampas  are  also 
found  in  the  caverns  of  Brazil,  where  the  climate  is  now,  and 


122  THE    NEOG^IC   REALM.  [CHAP. 

probably  always  has  been,  tropical.  Up  to  the  present,  it  is, 
accordingly,  impossible  to  account  satisfactorily  for  the  disappear- 
ance of  all  the  larger  forms  from  among  the  mammalian  fauna  of 
South  America. 

Returning  to  the  fauna  itself,  it  may  be  asserted  that  before 
the  great  intrusion  of  northern  forms  the  mammals  of  Neogaea 
were  far  more  distinct  from  those  of  the  rest  of  the  world  than 
is  the  case  with  the  fauna  of  any  other  region,  with  the  exception 
of  Australasia;  and  that  consequently  there  can  be  no  hesita- 
tion in  allowing  this  part  of  the  earth's  surface  to  take  rank  as  a 
primary  realm.  At  the  time  when  the  Santa  Cruz  fauna  was  so 
decidedly  marked  off  there  was  a  much  more  general  similarity 
between  the  faunas  of  all  the  other  regions  of  the  world  (exclusive 
of  Notogaea)  than  is  the  case  at  the  present  day,  and  it  is  this 
antiquity  of  the  differentiation  of  the  Neogaeic  fauna  that  supports 
so  strongly  its  claim  to  distinctness. 

It  has  been  suggested  that  the  first  land-connection  between 
South  and  North  America  was  probably  of  limited  extent  or  short 
duration  ;  and  some  evidence  of  a  later  separation  between  the 
two  areas  is  afforded  by  the  beds  of  marine  shells  already  mentioned 
as  occurring  in  the  upper  part  of  the  Pampean  deposits ;  these 
beds  marking  an  epoch  of  submergence  of  a  considerable  portion 
of  the  area1.  Subsequently  to  this  final  submergence  of  portions 
of  Argentina  and  Uruguay  there  was  a  great  upheaval  of  the 
country,  indicated  not  only  by  the  upraising  of  the  aforesaid  marine 
beds,  but  likewise  by  that  of  the  sand-dunes  fringing  most  parts  of 
the  Argentine  coast.  This  upheaval,  which  has  taken  place  within 
the  human  period,  certainly  resulted  in  the  final  union  of  the  two 
Americas ;  and  since  its  date  there  has  probably  continued  to  be 
a  greater  and  greater  admixture  of  the  two  originally  distinct 
faunas,  so  far  as  climatic  conditions  have  permitted.  It  is,  how- 
ever, not  a  little  curious  that  some  of  the  original  northern  types, 
such  as  the  vicunas  and  guanacos,  have  entirely  died  out  in  their 
original  habitat,  to  flourish  only  in  the  southern  half  of  the  con- 
tinent. 

1  There  is  some  evidence  to  show  that  the  isthmus  of  Panama  was  never 
completely  submerged  after  the  Pliocene,  but  it  may  have  been  so  narrow  as 
not  to  allow  of  much  migration  of  the  fauna. 


III.]  DISTINCTNESS   OF   EXISTING   FAUNA.  123 

Hitherto  especial  stress  has  been  laid  on  the  fossil  mammals 
of  Neogaea  as  entitling  the  tract  to  form  a  primary  Distinctness 
realm,  on  account  of  the  distinctness  of  its  fauna  of  the  existing 
from  that  of  the  rest  of  the  world  during  a  consider- 
able portion  of  the  Tertiary  epoch.  Even  at  the  present  day, 
however,  when,  as  already  shown,  its  mammalian  fauna  contains  a 
very  large  admixture  of  types  which  have  immigrated  from  the 
north  at  a  comparatively  recent  epoch,  it  still  stands  widely  apart 
from  other  regions.  On  this  point  may  be  quoted  the  admirable 
summary  given  in  "Island  Life"1  by  Dr  Wallace,  who  writes  that 
among  the  peculiar  mammals  we  have  "  the  prehensile-tailed 
monkeys  and  the  marmosets,  the  blood-sucking  bats,  the  coati- 
mundis,  the  peccaries,  the  llamas  and  alpacas  [vicunas  and 
guanacos],  chinchillas,  the  agutis,  the  sloths,  the  armadillos,  and 
the  ant-eaters ;  a  series  of  types  more  varied  and  more  distinct 
from  those  of  the  rest  of  the  world  than  any  other  continent  can 
boast  of.  Among  birds  we  have  the  charming  sugar-birds,  forming 
the  family  C&rebidce,  the  immense  and  wonderfully  varied  group 
of  tanagers  (Tanagridce),  the  exquisite  little  manakins  and  the 
gorgeously-coloured  chatterers  ( Cotingidce)  ;  the  host  of  tree- 
creepers  of  the  family  Dendrocolaptidce,  the  wonderful  toucans 
(Rhamphastid<z\  the  puff-birds  (Bucconidcz),  jacamars  (Galbu- 
lid<z),  todies  ( Todidce),  and  motmots  (Momotida) ;  the  marvellous 
assemblage  of  four  hundred  distinct  kinds  of  humming-birds 
(Trochilidce\  the  gorgeous  macaws  (Ara),  the  curassows  (Cracida), 
the  trumpeters  (Psophiidcf),  and  the  sun-bitterns  (Eurypygida). 
Here  again  there  is  no  other  continent  or  region  that  can  produce 
such  an  assemblage  of  remarkable  and  perfectly  distinct  groups  of 
birds ;  and  no  less  wonderful  is  its  richness  in  species,  since  these 
fully  equal,  if  they  do  not  surpass,  those  of  the  two  great  tropical 
regions  of  the  Eastern  Hemisphere  (the  Ethiopian  and  the 
Oriental)  combined."  Not  less  noteworthy  among  the  birds  are 
the  screamers  (PalctTiiedeidcz) ;  the  tinamus  ( Tinamida),  which 
while  outwardly  resembling  game-birds,  agree  with  the  struthious 
birds  in  the  structure  of  their  skulls ;  and  the  rheas  (Rheida),  or 
South  American  ostriches,  whose  nearest  allies  are  the  true 

1  Pages  50,  51.    In  this  quotation  the  scientific  names  of  some  of  the  groups 
have  been  added. 


124  THE    NEOG^IC    REALM.  [CHAP. 

ostriches  of  the  Old  World.  The  hoatzin  (Opisthocomus\  the  oil- 
bird  (Steatornis\  and  the  boat-bill  (Cancroma)  are  likewise 
peculiar  Neogaeic  types.  Still  more  remarkable  is  the  solitary 
Andean  survival  (Ccenolestes]  of  the  diprotodont  marsupials  of 
the  Santa  Cruz  epoch.  A  curious  feature  of  the  Neogaean 
forest-mammals — whether  they  belong  to  the  old  fauna  or  the 
new — is  the  frequency  of  prehensile  power  in  the  tail.  Not  only 
does  this  occur  in  several  genera  of  monkeys,  but  also  in  Cerco- 
leptes,  Synetheres,  Chcetomys,  Capromys  prehensilis,  Cydoturus, 
Didelphys,  and  Ccenolestes.  A  parallel  is  to  be  found  elsewhere 
only  in  Australia. 

After  referring  to  the  deficiency  of  the  many  types  of 
mammals  alluded  to  in  an  earlier  paragraph  of  the  present 
chapter,  the  author  adds  that  "Among  birds  we  have  to  notice  the 
absence  of  tits,  true  flycatchers,  shrikes,  sun-birds,  starlings,  larks 
(except  a  solitary  species  in  the  Andes),  rollers,  bee-eaters,  and 
pheasants,  while  warblers  are  very  scarce,  and  the  almost  cosmo- 
politan wagtails  are  represented  by  a  single  species  of  pipit 

Whether,  therefore,  we  consider  its  richness  in  peculiar  forms  of 
animal  life,  its  enormous  variety  of  species,  its  numerous  defici- 
encies as  compared  with  other  parts  of  the  world,  or  the  prevalence 
of  a  low  type  of  organisation  among  its  higher  animals,  the 
Neotropical  region  stands  out  as  undoubtedly  the  most  remark- 
able of  the  great  zoological  regions  of  the  earth." 

The  distinctness  is,  however,  by  no  means  confined  to 
mammals  and  birds.  Of  the  land  molluscs,  Mr  A.  H.  Cooke1 
writes  that  they  present  a  marked  contrast  to  those  of  North 
America.  "Instead  of  being  scanty,  they  are  exceedingly 
abundant;  instead  of  being  small  and  obscure,  they  are  among 
the  largest  in  size,  most  brilliant  in  colour,  and  most  singular 
in  shape  that  are  known  to  exist.  At  the  same  time  they 
are,  as  a  whole,  isolated  in  type,  and  exhibit  but  little  relation 
with  the  Mollusca  of  any  other  region." 

Having  arrived  at  the  conclusion  that  the  original  Neogseic 
mammalian  fauna,  exemplified  in  the  Santa  Cruz 

Origin  of  the 

Santa  Cruz          beds,  has  not  been  derived  from  North  America, 
it  remains  to  endeavour  to  account  for  its  origin. 
1   The  Cambridge  Natural  History — Mollusca,  p.  342  (1895). 


III.]  ORIGIN    OF   SANTA   CRUZ   FAUNA.  125 

This,  however,  is  a  difficult  and  perplexing  subject  which  it  is 
scarcely  possible  to  explain  fully  in  the  present  imperfect  state  of 
palaeontological  knowledge. 

With  regard  to  the  marsupials  of  an  Australian  type,  it  has 
already  been  stated  in  the  preceding  chapter1  that  these  appear  to 
have  been  derived  from  Notogaea  by  means  of  a  southern  land- 
bridge.  The  hypothesis  there  suggested  is  that  the  immigration 
has  taken  place  via  the  Antarctic  continent,  probably  across  the 
southern  Pacific2.  It  is  known  that  shallow  water  extends  from 
southern  Patagonia  andTierra  del  Fuego  to  South  Shetland;  while 
between  Australia  and  the  Antarctic  land  there  are  no  depths  ex- 
ceeding 2000  fathoms.  On  the  other  hand  it  is  just  possible  that 
the  connection  may  have  been  by  way  of  Polynesia. 

In  this  place  reference  may  be  made  to  certain  very  remark- 
able resemblances  existing  between  animals  of  groups  other  than 
mammals  respectively  inhabiting  Neogsea  and  Notogsea.  The  first 
instance  is  that  of  two  peculiar  families  of  freshwater  fishes,  known 
as  the  Haplochitonida  and  Galaxiidce.  Of  these  the  former  has 
one  Australian  and  a  second  South  American  genus,  while  the 
latter  is  represented  by  a  single  genus  (Galaxias),  common  to 
New  Zealand,  Australia,  and  the  extremity  of  South  America. 
This,  however,  is  by  no  means  all,  since  one  species  of  the  last- 
mentioned  genus  (G.  attenuatus)  is  found  in  regions  as  remote  from 
one  another  as  New  Zealand  and  Tasmania  on  the  one  hand,  and  the 
Falkland  Islands  and  the  extremity  of  Patagonia  on  the  other. 
Commenting  on  this,  Dr  Wallace  remarks3,  it  is  impossible  to 
believe  that  a  land  connection  between  South  America  and 
Notogaea  could  have  existed  "  within  the  period  of  existence  of 
this  one  species  of  fish,  not  only  on  account  of  what  we  know  of 
the  permanency  of  continents  and  deep  oceans ;  but  because  such 
a  connection  must  have  led  to  much  more  numerous  and  im- 
portant cases  of  similarity  of  natural  productions  than  we  actually 
find.  Rather  must  we  look  to  the  transport  of  the  ova  across  the 

1  Supra,  p.  55. 

2  Possibly,  as  suggested  below,  the  connection  may  have  been  nearer  the 
tropics. 

3  Geographical  Distribution  of  Animals,  Vol.  I.  pp.  401,  402;  see  also  Vol. 
II.  pp.  82,  83. 


126  THE   NEOG^IC   REALM.  [CHAP. 

southern  seas,  aided  perhaps  by  the  Antarctic  ice,  and  a  former 
greater  extension  of  South  America  towards  the  pole."  After 
remarking  how  such  transmission  might  take  place  with  but  little 
extension  of  the  present  Antarctic  lands,  Dr  Wallace  adds  that 
"  there  is  evidently  some  means  by  which  ova  or  young  fishes  are 
carried  moderate  distances,  from  the  fact  that  remote  Alpine  lakes 
and  distinct  river-systems  often  have  the  same  species.  Glaciers 
and  icebergs  generally  have  pools  of  fresh  water  on  their  surfaces ; 
and  whatever  cause  transmits  fish  to  an  isolated  pond  might 
occasionally  stock  these  pools,  and  by  this  means  introduce  the 
fishes  of  one  southern  island  into  another." 

Allowing  all  due  weight  to  these  objections  to  a  land  con- 
nection between  Notogaea  and  Neogsea,  it  seems  almost  im- 
possible to  believe  that  the  transit  has  taken  place  in  the  manner 
suggested  by  Dr  Wallace. 

Another  piece  of  evidence  is  afforded  by  some  observations  of 
Mr  F.  E.  Beddard l  in  regard  to  the  intimate  relationship  existing 
between  the  earth-worms  of  New  Zealand  and  Eastern  Australia 
on  the  one  hand,  and  those  of  Patagonia  on  the  other.  Without 
committing  himself  to  any  theory  as  to  how  the  communication 
took  place,  the  author  is  content  to  say  that  "the  facts  seem  to 
point  to  a  more  recent  communication  between  Patagonia  and 
New  Zealand  than  between  either  of  those  countries  and  the  Cape 
of  Good  Hope." 

Assuming  a  land  connection,  earth-worms  would  suggest  that 
it  was  probably  not  in  very  high  latitudes.  Now  I  have  been 
informed  on  verbal  authority  that  there  is  a  curious  similarity 
between  the  slugs  of  Patagonia  and  those  of  Polynesia.  And  it  is 
probable  that  the  latter  tract  indicates  a  subsiding  area  which 
was  formerly  connected  with  Patagonia.  Possibly,  therefore,  there 
may  have  been  a  land  connection  between  Patagonia  and  Australia 
via  Polynesia ;  and  this  may  have  been  the  line  through 
which  Neogsea  received  the  Notogseic  elements  in  its  fauna. 
Whether  it  could  have  existed  at  a  date  sufficiently  late  for  the 
passage  of  the  marsupials,  it  is  impossible  to  say.  If  it  existed, 
it  probably  allowed  only  a  limited  communication  between  the 

1  Appendix,  No.   5,  pp.  170,  171. 


III.]  ORIGIN   OF   SANTA   CRUZ   FAUNA.  I2/ 

Notogaeic  and  Neogaeic  mammals ;  and  it  is  easy  to  imagine  that 
the  Polynesian  mammals  (if  they  existed)  were  drowned  out  by 
submergence,  as  has  undoubtedly  been  the  case  with  many  of 
those  of  the  West  Indies.  In  dismissing  this  part  of  the  subject, 
it  may  be  observed  that  it  appears  impossible  to  adequately 
explain  the  presence  of  a  Notogaeic  element  in  the  fauna  of 
Neogaea  without  the  aid  of  some  form  of  southern  land  connection; 
although  there  is  not  sufficient  evidence  to  show  in  what  latitude 
such  connection  (or  connections)  existed. 

Attention  must  now  be  directed  to  the  Santa  Crucian  mammals 
other  than  marsupials.  With  the  exception  of  the  edentates,  which 
probably  originated  in  Neogsea,  or  possibly  in  some  still  more 
southern  land,  all  the  evidence  points  to  the  whole  of  these 
being  originally  of  northern  derivation ;  the  ungulates  having 
affinities  with  the  ancestral  types  from  which  the  earlier  Tertiary 
perissodactyles  were  descended,  while  the  rodents  have  certain 
relationships  with  the  early  European  members  of  the  order.  The 
monkeys  again  were  probably  descended  from  lemuroids ;  and 
the  solenodons  are  evidently  related  to  the  Old  World  insectivores. 
It  has  been  shown  that  this  portion  of  the  fauna  did  not  come 
from  North  America;  and  it  is  certainly  not  derived  from  Notogaea. 
Accordingly,  the  only  route  by  which  it  could  have  entered  is  by 
way  of  Africa.  The  only  marked  community  between  the  Ethio- 
pian and  Neogaeic  faunas  as  regards  mammals  relates  to  the  hystri- 
comorphous  rodents ;  but  this  community  is  a  very  marked  one, 
and  difficult  to  explain  on  any  other  hypothesis  than  that  of  a 
connection  between  the  two  areas.  The  possibility  of  a  close 
community  of  origin  between  the  toxodonts  and  the  hyraces  has 
already  been  mentioned  ;  and  if  it  be  substantiated,  it  will  be 
highly  important.  Of  course,  on  the  supposition  of  an  African 
origin  for  the  eutherian  mammalian  fauna  of  Tertiary  Neogaea, 
it  must  be  taken  for  granted  that  the  ancestral  types  entered 
Africa  long  before  the  progenitors  of  its  modern  fauna  ; 
although  probably  not  before  the  ancestors  of  the  Malagasy 
fauna.  It  may  be  objected  that  we  ought  to  find  Neogaeic 
Tertiary  types  of  ungulates  in  Africa  ;  but  we  are  unacquainted 
with  the  Tertiary  palaeontology  of  that  country,  and  it  is 
quite  probable  that  the  peculiar  subordinal  Neogaeic  types  of 


128  THE    NEOG^IC    REALM.  [CHAP. 

ungulates  were  only  developed  as  such  in  America  itself.  Even 
if  they  ever  existed  in  Africa  there  is  no  more  reason  why  they 
should  have  survived  there  than  in  America.  As  the  evidence 
for  the  presence  of  Insectivora  in  the  Santa  Cruz  deposits  is 
not  very  strong,  the  case  of  the  West  Indian  solenodons  must 
not  be  pressed  too  strongly,  but  their  affinity  to  the  tenrecs  of 
Madagascar,  and  the  absence  of  allied  types  in  the  North  American 
Tertiaries,  both  point  to  their  having  reached  Neogsea  with  the 
other  eutherians. 

Regarding  a  possible  connection  between  Africa  and  South 
America  by  way  of  the  Antarctic  continent,  Dr  Blanford1  writes 
as  follows:  —  "Singularly  enough,  so  far  as  our  present  information 
as  to  the  depths  of  the  southern  oceans  goes,  there  would  appear 
at  first  sight  to  be  less  difficulty  in  supposing  a  former  extension 
of  the  southern  continent  to  Australia  and  South  America  than  to 
Africa,  the  depth  as  shown  on  the  '  Challenger'  charts  south  of  the 
former  continents  nowhere  exceeding  2000  fathoms,  whereas  to  the 
south  of  Africa  there  is  represented  a  considerable  belt  of  greater 
depth.  But  on  an  Admiralty  chart  on  which  all  the  known  deep 
soundings  are  marked,  none  are  shown  south  of  the  southern 
extremity  of  Africa....  So  far  as  our  present  information  goes, 
the  ocean  south  of  the  Cape  of  Good  Hope  may  be  no  deeper 
than  the  Mozambique  channel,  though  probably  the  depth  is 
greater  in  the  former  case." 

Before  discussing  certain  relationships  between  the  Ethiopian 
fauna  and  that  of  Neogaea,  it  seems  advisable  to 

"Antarctica"  .  .  .  _ 

and  the  South      refer  to  some  recent  views  as  to  the   existence  of 


a  Sreat  southern  circumpolar  continent  in  Tertiary- 
Ethiopian  times,  extending  into  comparatively  low  latitudes, 

and  connected,  at  all  events  temporarily,  with 
America,  Africa,  and  Australia.  For  this  continent  the  name 
Antarctica  has  been  suggested  by  Dr  H.  O.  Forbes2,  who  urges 
that  many  of  the  types  of  animal  life  now  confined  to  the  southern 
hemisphere  have  originated  there.  It  is  chiefly  to  show  the  fallacy 
of  these  latter  views  that  the  subject  is  referred  to  here  ;  palaeon- 
tological  evidence  clearly  proving  that  several  of  the  groups  of 

1  Appendix,  No.  8,  p.  100. 

2  Appendix,  No.  15. 


III.]         NORTHERN   ORIGIN   OF   SOUTHERN   FORMS.          1 29 

animals  assumed  to  be  essentially  southern,  really  had  a  northern 
origin.  It  may  be  premised  that,  according  to  the  view  of 
Dr  Forbes,  "  Antarctica  "  followed  nearly  the  2000  fathom  line, 
extending  northwards  from  a  circumpolar  area  by  broad  expan- 
sions, one  to  join  an  old  New  Zealand  continental  island  (includ- 
ing the  Antipodes,  Macquarries,  New  Zealand,  and  Chatham,  Lord 
Howe,  Norfolk,  and  the  Kermadec  and  Fiji  Islands);  a  second  to 
East  Australia  and  Tasmania ;  a  third  to  the  Mascarene  and 
adjacent  islands  ;  perhaps  one  to  South  Africa ;  and  finally  one 
to  South  America. 

As  regards  the  marsupials,  which  are  among  those  considered 
to  be  southern  types,  the  evidence  of  the  northern  Jurassic  and 
Cretaceous  kinds  alluded  to  in  the  preceding  chapter,  coupled 
with  the  presence  of  opossums  in  the  Oligocene  of  the  northern 
hemisphere,  renders  it  practically  certain  that  the  group  did  not 
originate  in  the  southern  hemisphere. 

Among  other  groups  cited  by  Dr  Forbes  as  being  mainly  or 
exclusively  southern  in  their  distribution  and  origin  are  the  parrots 
(Psitiaci)  and  trogons  (Trogonidce).  But  both  these  are  represented 
in  a  fossil  state  in  the  Oligocene  strata  of  France,  and  are  thus 
shown  to  have  been  originally  denizens  of  the  temperate  regions 
of  the  northern  hemisphere.  Take,  again,  the  case  of  the  struthious 
birds,  or  Ratitse,  which  although  cited  as  a  southern  group,  are 
represented  by  an  ostrich  (Struthio)  in  the  Pliocene  of  Northern 
India  and  the  Crimea,  while  the  former  deposits  have  yielded 
remains  of  a  three-toed  genus  allied  probably  to  the  emeus  and 
cassowaries.  Much  the  same  may  be  said  in  regard  to  the  giant 
land-tortoises  (Testudo\  which  although  now  confined  to  the 
Galapagos  and  Mascarene  islands,  were  abundant  in  Northern 
India,  Greece,  France,  and  the  United  States  during  the  Pliocene, 
and  also  occur  in  the  French  Miocene  and  Oligocene,  as  well  as 
in  the  Plistocene  deposits  of  the  Maltese  caves.  The  group  was 
thus  evidently  a  northern  one  originally,  and  as  it  is  unknown  in 
the  southern  hemisphere  before  the  Pampean  epoch  of  Argentina 
and  the  superficial  deposits  of  Madagascar,  its  southern  migration 
probably  did  not  take  place  till  the  Miocene,  or  even  the 
Pliocene.  Indeed,  the  separation  of  North  and  South  America 
indicates  that,  if  the  Galapagos  tortoises  came  from  the  former 
L.  9 


130  THE    NEOG^EIC   REALM.  [CHAP. 

country,  they  could  not  have  reached  their  present  habitat  till  the 
end  of  the  Miocene. 

On  the  question  of  the  southern  or  northern  origin  of  some  of 
the  above-mentioned  birds,  Professor  Huxley1,  so  far  back  as  1868, 
wrote  as  follows: — "I  watch  the  progress  of  M.  Alphonse  Milne- 
Edwards's  researches  with  great  interest,  to  know  whether  parrots, 
pigeons,  Dromceidce  (Casuariidas)  and  Rhtzidce  occur  in  force,  or 
at  all,  among  the  Miocene  birds.  If  they  are  absent  from  the 
Miocene  fauna  of  Arctogaea,  it  will  be  necessary  to  suppose  that 
these  groups  of  birds  are  of  sufficiently  ancient  origin  to  have  been 
separated,  even  before  the  Miocene  epoch  in  Austro-Columbia 
(Neogaea)  and  Australasia,  whence  they  have  subsequently 
colonised  part  of  Arctogsea ;  while,  on  the  other  hand,  their 
presence  in  European  Miocene  formations  will  render  it  possible 
that  the  colonisation  has  taken  place  the  other  way,  and  that  these 
birds  have  attained  their  wonderful  multiplicity  and  diversity  of 
forms  in  Austro-Columbia  and  Australasia  simply  in  consequence 
of  the  very  favourable  nature  of  the  conditions  to  which  they  have 
been  exposed  in  that  country. 

"I  confess  I  incline  to  the  latter  supposition.  The  distribution 
of  Psittacula,  for  instance,  is  quite  unintelligible  to  me  upon  any 
other  supposition  than  that  this  genus  existed  in  the  Miocene 
epoch,  or  earlier,  in  Northern  Arctogaea,  and  has  thence  spread 
into  Austro-Columbia,  South  Africa,  India,  and  the  Papuan 
Islands,  where  it  is  now  found." 

Although  the  term  Psittacula  has  now  been  restricted  so  as  to 
include  only  the  Neogaeic  forms,  this  passage  is  almost  prophetic ; 
both  parrots  and  pigeons  having,  as  already  stated,  been  dis- 
covered in  the  French  Oligocene,  while  the  Australian  and 
probably  the  South  American  ratite  birds  appear  to  have  had  an 
Indian  forerunner.  And  here  it  may  be  mentioned  that  the 
South  American  ostriches  (Rhea]  which  are  primitive  types  allied 
to  the  ostrich,  would  seem  to  have  made  their  way  into  Neogaea 
via  Africa,  as  there  are  no  traces  of  ancestral  forms  in  the  North 
American  Tertiaries. 

On  the  other  hand,  there  is  considerable  probability  that  the 

1  Appendix,  No.   18,  p.  319. 


III.]          NORTHERN    ORIGIN   OF   SOUTHERN   FORMS.          131 

penguins  (Spheniscidcz),  which  present  a  relation  to  other  birds 
somewhat  analogous  to  that  exhibited  by  the  edentates  to  other 
mammals,  having  no  apparent  affinity  with  any  group — -may  prove 
to  be  an  exception  to  the  rule  of  the  northern  origin  of  most  of  the 
existing  southern  types  of  terrestrial  vertebrates,  since  they  are 
quite  unknown  in  the  north,  and  occur  fossil  both  in  New  Zealand 
and  Patagonia.  ',  / 

Another  marked  instance  of  the  northern  origin  of  southern 
types  is  afforded  by  the  side-necked,  or  pleurodiran  Chelonia, 
which  although  now  restricted  to  the  more  southern  parts  of  the 
globe,  were  abundant  during  Secondary  and  early  Tertiary  times 
throughout  the  northern  hemisphere.  A  striking  example  of  this 
is  shown  in  the  family  Pelomednsida,  whose  existing  representatives 
are  confined  to  Africa,  Madagascar,  and  South  America.  Among 
these,  two  out  of  three  genera,  namely  Sternothoerus  and  Pelomedusa 
are  found  in  Ethiopian  Africa  and  Madagascar,  one  of  them  also 
ranging  into  the  Sinai  tic  peninsula ;  while  the  third  (Podocnemis} 
has  five  species  in  South  America  and  a  sixth  in  Madagascar. 
There  occurs,  however,  in  the  upper  Cretaceous  of  the  United 
States  the  allied  extinct  genus  Bothremys,  and  Podocnemis  itself  is 
represented  in  the  London  Clay  and  the  Eocene  of  the  Punjab. 
Here  the  inference  would  seem  to  be  that  the  latter  genus 
originated  in  the  northern  half  of  the  Old  World,  passed  by  way  of 
India  into  Madagascar  and  Africa,  and  thence  by  a  southern  route 
into  Neoggea.  Even  if  this  particular  genus  occurred  in  the  early 
Eocene  of  North  America,  which  it  does  not,  it  could  scarcely  have 
crossed  the  sea  into  South  America ;  and  the  migration  can  hardly 
have  taken  place  since  the  union  of  the  two  continents.  In 
commenting  on  the  distribution  of  Podocnemis  Dr  Blanford1 
observes  that  as  the  incursion  of  more  modern  types  into  Africa 
appears  to  have  driven  out  many  of  the  older,  it  is  in  Madagascar 
that  traces  of  the  relationship  of  the  modern  fauna  to  that  of 
Neogaea  should  be  looked  for.  One  such  instance  is  the  occur- 
rence there  of  Podocnemis^  and  a  second  that  of  the  Centetidcz  in 
that  island.  Perhaps  the  occurrence  of  sucker-footed  bats  only  in 
Brazil  where  they  are  represented  by  the  single  species  of 

1  Appendix,  No.  8,  p.  101. 

9—2 


132  THE    NEOG^IC   REALM.  [CHAP. 

Thyropoda,  and  in  Madagascar,  where  there  is  the  sole  member  of 
the  allied  genus  Myxopoda^  may  be  an  analogous  instance. 

In  the  second  family  of  the  pleurodiran  Chelonia,  the  Chelyidcz, 
which  are  now  restricted  to  South  America  and  Australia  (the 
genera  in  the  two  areas  being  distinct),  there  is  at  present  no 
evidence  of  the  derivation  of  the  Australian  forms,  but  of  the 
Neogaeic  types  Platcmys  is  represented  in  the  Cretaceous  of  the 
United  States,  and  Hydraspis  in  the  Eocene  of  Bombay.  Although 
the  northern  origin  of  the  family  is  thus  proved,  the  explanation 
of  how  the  existing  forms  attained  their  present  distribution  is 
very  difficult.  Possibly  Hydraspis  may  have  reached  South 
America  by  way  of  Africa;  but  it  is  difficult  to  believe,  in  the 
absence  of  its  remains,  that  Platemys  survived  in  North  America 
till  the  late  Miocene  communication  with  Neogaea  was  estab- 
lished. 

Among  snakes,  the  boas  of  the  genera  Corallus  and  Boa  are 
confined  to  South  America  and  Madagascar,  and  thus  have 
precisely  the  same  distribution  as  Podocnemis.  Now  although  true 
boas  are  unknown  as  fossils,  the  allied  extinct  genus  Paleryx 
occurs  in  the  European  Oligocene,  thus  pointing  to  the  northern 
origin  of  the  group,  which  has  probably  reached  South  America  by 
way  of  Madagascar  and  Africa. 

Another  remarkable  case  is  afforded  by  the  limbless  lizards  of 
the  family  Amphisbcenidcz,  which  are  now  almost  equally  divided 
between  South  America  and  South  Africa,  although  one  genus 
extends  into  the  Mediterranean  area,  and  two  are  found  in  North 
America ;  the  two  genera  Amphisb&na  and  Anops  being  common 
to  South  America  and  Africa,  while  the  northern  ones  are  different. 
The  northern  origin  of  the  family  is,  however,  indicated  by  the  re- 
cent discovery  of  fossil  forms1  in  the  White  River  Oligocene  of  the 
United  States.  Here  the  evidence  strongly  points  to  a  southern 
connection  between  Neogaea  and  Africa;  Tertiary  forms  having 
probably  existed  in  Europe  or  Asia  as  well  as  in  North  America. 
A  second  instance  that  may  be  cited  among  lizards  is  the  family 
of  the  Iguanidcz,  which  while  now  mainly  Neogaeic,  has  represen- 
tatives in  the  warmer  parts  of  North  America,  and  also  includes 

1  Baur,  American  Naturalist,  Vol.  xxvii.  p.  998  (1893). 


III.]  CONNECTION   WITH   AFRICA.  133 

two  outlying  genera  in  Madagascar,  and  a  third  in  the  Fiji 
and  Friendly  Islands.  But  fossil  iguanas  occur  in  the  French 
Oligocene,  and  it  may  hence  be  suggested  that  the  group  may 
have  reached  Neogaea  via  Madagascar  and  Africa;  while  if 
the  connection  between  Patagonia  and  Polynesia  alluded  to  above 
were  substantiated,  the  origin  of  the  Polynesian  forms  could  be 
accounted  for. 

During  the  middle  portion  of  the  Secondary  period  a  very 
curious  resemblance  between  the  fauna  of  Ethiopia  and  Neogaea 
is  exhibited  by  the  occurrence  in  both  of  certain  very  peculiar 
reptiles  known  as  Mesosaurns  (Stereosternum),  which  have  been 
referred  to  the  Sauropterygia.  Remains  of  these  reptiles  have 
been  obtained  at  San  Paolo  in  Brazil,  and  in  Griqualand  West 
and  other  parts  of  South  Africa,  but  nowhere  else ;  and,  although 
the  type  may  be  of  northern  origin,  this  curious  distribution 
apparently  points  strongly  to  a  connection  between  Africa  and 
South  America  as  far  back  as  the  Secondary  epoch.  This  con- 
nection, as  pointed  out  by  Neumayr1,  was,  however,  probably  by 
way  of  the  Atlantic. 

Somewhat  similar  relationships  to  those  of  living  reptiles  are 
exhibited  by  fishes,  among  which  the  ffaplochitonidce  and 
Galaxiida  have  been  already  mentioned.  Very  remarkable  is  the 
case  of  the  lung-fishes  Lepidosirenidce,  where  there  is  a  very  close 
relationship  between  the  West  African  Protopterus  and  Lepidosiren 
of  Brazil  and  Paraguay  ;  the  Australian  Ceratodus  being  markedly 
distinct  from  both.  Although  the  two  former  are  unknown  as 
fossils,  teeth  of  the  latter  are  abundant  in  the  Trias  and  Jurassic  of 
Europe,  India,  South  Africa,  and  the  United  States ;  while  during 
the  Palaeozoic  era  extinct  families  of  the  subclass  (Dipnoi)  were 
abundant  in  the  northern  hemisphere.  Clearly,  then,  the  group  was 
originally  northern  in  origin;  and  Ceratodus  apparently  migrated 
south  both  into  Africa  and  Australia.  Taking  into  account  the 
Cretaceous  separation  of  North  and  South  America,  and  the  close 
alliance  between  Lepidosiren  and  Protopterus,  it  is,  however, 
difficult  to  see  how  the  latter  reached  its  present  habitat  except  by 
way  of  Africa.  If  this  be  so,  and  the  connection  between  South 

1   Vide  supra,  p.  IT 8,  note. 


134  THE    NEOG^EIC   REALM.  [CHAP. 

Africa  and  South  America  in  Tertiary  times  was  only  in  high 
latitudes,  a  warm  epoch  in  the  southern  hemisphere  must  have 
been  necessary  for  the  passage  of  such  tropical  forms.  It  might 
be  urged  that  as  Ceratodus  dates  from  the  Trias,  the  other  two 
genera  might  have  reached  their  present  habitats  at  a  very 
distant  epoch ;  but  their  specialisation  is  against  their  antiquity. 
Another  family  which  is  essentially  southern  is  that  of  the  Osteo- 
glossidce,  represented  by  Arapaima  of  the  Brazilian  rivers,  and 
Osteoglossum,  with  one  species  from  Brazil  and  the  Guianas,  a 
second  from  Sumatra,  and  two  others  from  Australia.  But  the 
northern  origin  of  the  family  is  indicated  by  the  occurrence  of  the 
extinct  genus  Dapedoglossus  in  the  Eocene  of  Wyoming.  Here 
there  is  a  presumption  that  Osteoglossum  originated  in  Asia,  from 
which  it  passed  in  one  direction  by  way  of  Malaysia  to  Australia, 
and  in  another  through  Africa  to  South  America.  Two  other 
families  of  freshwater  fishes  have  a  somewhat  similar  distribution ; 
the  first  being  the  Chromidid<zy  which  includes  spiny  fishes  mainly 
characteristic  of  tropical  America  and  Africa,  but  extending 
eastwards  into  Syria,  and  sparingly  represented  in  Southern  India 
and  Ceylon.  In  a  fossil  state  they  occur  in  the  Cretaceous  of 
Syria ;  and,  although  none  of  the  genera  are  common  to  the  two 
continents,  they  are  highly  suggestive  of  a  connection  between 
Africa  and  South  America.  The  second  family  is  that  of  the 
Characiniidce,  comprising  fish  more  nearly  allied  to  the  carps,  and 
now  exclusively  confined  to  tropical  America  and  Africa.  Although 
the  palaeontological  record  is  a  blank,  this  can  scarcely  be  taken 
as  a  sufficient  indication  that  the  family  has  always  been  a 
southern  one. 

From  the  foregoing  facts  it  may  be  considered  that  the  as- 
sumption of  an  Antarctic  continent  is  unnecessary 
to  explain  the  origin  of  the  many  forms  of  vertebrate 
life  which  are  now  exclusively  or  mainly  southern  ;  nearly  all  of 
these,  with  the  exception  of  the  edentates  and  penguins,  being  of 
northern   derivation,    and   thus    apparently  showing   a   southern 
migration  of  the  older  forms  of  life.     The  Cretaceous  and  Tertiary 
break  between   North  and  South  America  appears,  however,  to 
have  prevented  the  occurrence  of  such  migration  in  the  western 
hemisphere  till  the  close  of  the  Miocene :  and  it  is  accordingly 


III.]  SUB-REGIONS.  135 

necessary  to  look  elsewhere  for  the  origin  of  the  Neogaeic  fauna. 
That  Africa  has  been  the  great  feeder  appears  the  most  probable 
explanation  ;  although  in  the  case  of  the  marsupials  it  seems 
necessary  to  look  to  Notogaea  as  the  point  of  origin.  Clearly, 
however,  the  presumed  connections  between  Neogaea,  Notogsea, 
and  Africa  have  not  been  very  continuous  or-  very  extensive 
in  Tertiary  times,  or  the  faunas  of  these  areas  would  have  been 
more  alike  than  they  are ;  and  this  suggests  that  the  northern 
extension  of  Antarctica  has  not  been  so  great  as  has  been 
supposed.  Whether  the  presumed  connection  between  Notogaea 
and  Neogaea  has  taken  place  by  way  of  Antarctica  or  Polynesia 
may  be  left  an  open  question.  With  regard  to  Africa,  the  recent 
researches  of  Dr  Gregory1  on  the  West  Indian  corals,  in  the 
course  of  which  it  is  urged  that  a  shallow-water  connection  "ex- 
tended across  the  Central  Atlantic  in — at  latest — Miocene  times," 
while  the  southward  extension  of  the  Atlantic  is  a  comparatively 
recent  feature,  indicate  the  possibility  that  the  land-connection 
which  existed  in  Jurassic  times  between  Brazil  and  Western 
Africa  may  have  persisted  till  the  Tertiary  era. 

As   already  mentioned,  the   Neogaeic   realm   includes  but   a 
single  region — the   Neotropical :    and  in   this  four 

,..,,..,,  ,  Sub-regions. 

sub-regions  have  been  defined,  and  are  named  as 
follows.  Firstly  we  have  the  Brazilian  sub-region,  which  includes 
not  only  Brazil,  but  likewise  the  Guianas,  Venezuela,  Colombia, 
Ecuador,  Paraguay,  and  those  portions  of  Peru  and  Bolivia  lying 
on  the  Brazilian  side  of  the  Andes,  together  with  the  eastern 
slopes  of  that  portion  of  the  great  mountain-chain  itself.  This  is 
essentially  an  area  of  dense  tropical  forests,  locally  interspersed 
with  open  pastures,  or  "campos."  The  second  is  the  Chilian 
sub-region,  comprising  Chili,  Argentina  proper,  Uruguay,  Pata- 
gonia, and  such  portions  of  Peru  and  Bolivia  as  are  not  included 
in  the  preceding.  It  is  chiefly  an  area  of  open  plains  and 
pampas,  although  including  the  high  Andes.  Thirdly,  there  is 
the  Mexican  sub-region,  which  embraces  the  isthmus  of  Panamk, 
Central  America,  and  Southern  Mexico,  and  may  be  regarded  to 
a  great  extent  as  a  transitional  tract  between  the  typical  Neo- 

1  Quart.  Journ.  GcoL  Soc.  Vol.  LI.  pp.  306 — 307  (1895). 


136  THE   NEOG^EIC    REALM.  [CHAP. 

tropical  and  the  Sonoran  regions.  Lastly,  the  Antillean  sub- 
region  includes  the  West  Indian  Islands,  exclusive  of  Trinidad, 
which  for  zoological  purposes  may  be  regarded  as  part  and  parcel 
of  the  South  American  continent. 

From  the  survey  of  the  fossil  forms  it  has  been  shown  that 
during  the  Plistocene  epoch  the  mammalian  fauna  of  the  Chilian 
and  Brazilian  sub-regions  was  similar,  and  it  may  consequently  be 
inferred  that  the  present  differentiation  of  the  two  areas  in  this 
respect  is  a  comparatively  modern  feature,  probably  due  to 
the  disappearance  of  the  forests  from  the  Argentine.  At  the 
present  time  the  mammalian  fauna  of  the  Brazilian  sub-region  is 
essentially  that  of  the  Neotropical  region  as  a  whole,  nearly  all  the 
characteristic  groups  being  present  within  its  limits,  while  several 
are  almost  or  quite  peculiar  to  it.  Among  the  latter,  the  great 
ant-eater  (Myrmecophaga)  is  practically  confined  to  this  sub-region  \ 
while  most  of  the  sloths  and  marmosets  are  limited  to  it,  although 
a  few  extend  northwards  into  or  through  the  isthmus.  The  pacas 
(Ccelogenys),  and  the  giant  armadillo  (Priodon}— the  sole  repre- 
sentative of  its  genus — are  likewise  restricted  to  this  tract ;  as 
is  the  bush-dog  (Icticyon),  and  also  one  genus  of  tree-porcupines 
( Chcetomys],  while  most  of  the  spiny  rats  (Echinomys  and  Loncheres) 
are  confined  to  it.  The  carpincho  (Ifydroc/icerus) — the  largest 
of  living  rodents  —  likewise  chiefly  pertains  to  the  Brazilian 
region,  although  extending  southwards  into  Uruguay.  The 
American  monkeys  are  also  very  abundantly  represented  here ; 
the  genera  Lagothrix,  Pithetia,  Brachyurus,  Brachy teles  (Eriodes), 
and  Callithrix  being  restricted  to  it.  Among  the  forms  that  are 
unrepresented,  may  be  mentioned  guanacos  and  vicunas  (Lama), 
viscachas  (Lagostomus),  the  Patagonian  cavy  (Dolichotis),  and 
chinchillas  (Eriomys  and  Lagidium). 

The  Mexican  or  Central  American  sub-region  differs  chiefly 
from  the  last  by  the  paucity  of  the  essentially  Neotropical  forms, 
and  the  large  mingling  of  Arctogaeic  types ;  among  the  latter, 
the  shrews  (Soriridce),  a  pouched  rat  (Heteromys}^  and  the  caxo- 
mistle  (Bassariscus)  being  noticeable.  In  the  Chilian  sub-region 
marmosets,  monkeys,  sloths,  tapirs,  and  peccaries  are  wanting; 

1  According  to  Senor  Figueira  it  just  enters  Uruguay. 


III.]  SUB-REGIONS.  137 

while  the  carpincho,  as  already  mentioned,  only  borders  on  it  in 
Uruguay.  Among  the  characteristic  types  are  prominent  the 
vicunas  of  the  Andes,  the  guanacos  of  the  Argentine  pampas  and 
Patagonia,  the  spectacled  bear  of  the  Andes,  the  chinchillas  of  the 
same  elevated  regions,  together  with  the  aquatic  coypu  (Myopo- 
tamus},  the  burrowing  viscacha,  and  the  cursorial  Patagonian 
cavy ;  all  the  three  latter  being  plain-dwelling  forms.  Armadillos 
are  abundant;  and  among  these  the  sub-family  represented  by  the 
beautiful  little  pichiciagos,  or  fairy-armadillos  (Chlamydophorus) 
is  peculiar,  one  of  the  two  species  inhabiting  open  plains  near 
Mendoza,  in  the  Argentine,  while  the  second  (regarded  by  some 
as  a  distinct  genus)  is  found  in  the  Bolivian  highlands. 

The  Antillean,  or  West  Indian  sub-region,  which  comprises  the 
West  Indian  Islands  (exclusive  of  Trinidad,  Tobago,  and  some  of 
the  adjacent  islets,  which  are  zoologically  a  part  of  continental 
South  America),  differs  widely  from  the  other  three  by  the  extreme 
poverty  of  its  mammalian  fauna;  monkeys,  marmosets,  carnivores, 
and  edentates  being  wanting,  and  the  class  mainly  represented  by 
bats,  insectivores,  and  rodents,  although  a  species  of  aguti  (Dasy- 
procta  antilliensis)  is  found  in  the  islands  of  St  Vincent  and  Santa 
Lucia,  in  the  Lesser  Antilles  group,  as  also  in  Tobago.  In 
addition  to  a  single  species  of  white-footed  mouse  (Sitomys)  said  to 
inhabit  Hayti  and  Martinique,  and  which  may  also  occur  in  some 
of  the  other  islands,  the  West  Indian  sub-region  is  especially 
characterised  by  the  large  arboreal  rodents  known  as  hutias,  which, 
while  belonging  to  the  family  Octodontidce,  represent  two  genera 
totally  unknown  elsewhere.  Of  these,  the  genus  Plagiodon  has 
but  a  single  species,  confined  to  Hayti  and  Jamaica;  although  in 
the  allied  Capromys  three  existing  species  are  found  in  Cuba,  and 
the  fourth  in  Jamaica,  the  extinct  kind  occurring  in  the  former 
island1.  The  nearest  relative  of  the  hutias  appears  to  be  the 
South  American  coypu,  but  the  group  seems  also  to  show  affini- 
ties with  the  porcupines.  From  caves  in  the  small  island  of 
Anguilla,  at  the  northern  extremity  of  the  Lesser  Antilles  group 
have  been  obtained  remains  of  a  large  extinct  beaver-like  rodent 
known  as  Amblyrhiza  (Loxomylus),  which  has  also  been  recorded 

1  Chapman,  Bull.  Amer.  Mus.  Vol.  IV.  p.  314  (1892). 


138  THE    NEOG^IC   REALM.  [CHAP. 

from  the  Pliocene  of  Argentina — a  fact  of  importance  as  serving 
to  connect  the  Antillean  fauna  with  that  of  the  mainland.  As 
mentioned  above,  this  genus  belongs  to  a  family  (Castoroididce) — 
typified  by  the  extinct  Castoroides  of  the  Plistocene  of  Ohio  and 
Georgia — with  species  which  rivalled  a  bear  in  point  of  size.  The 
other  Antillean  native  mammals  (exclusive  of  the  bats,  to  which  it 
will  be  unnecessary  to  refer)  are  the  two  species  of  the  genus 
Solenodon  respectively  inhabiting  Cuba  and  Hayti,  and  constitut- 
ing by  themselves  a  separate  family  among  the  Insectivora.  It 
has  been  already  mentioned  that  the  nearest  allies  of  these  strange 
creatures  are  the  tenrecs  (Centetidce)  of  Madagascar;  and  thus  both 
are  probably  derived  from  unknown  extinct  insectivores  formerly 
inhabiting  the  northern  hemisphere.  As  Jamaica  and  probably 
several  other  of  the  West  Indian  islands  contain  large  masses  of 
sedimentary  deposits  of  Tertiary  age,  it  is  probable  that  they 
come  under  the  denomination  of  continental  islands;  and  there 
seems  little  doubt,  from  the  evidence  of  their  mammals  alone,  that 
they  have  been  connected  with  the  mainland1.  Dr  Wallace  is  of 
opinion  that  "originally  they  probably  formed  part  of  Central 
America,  and  may  have  been  united  with  Yucatan  and  Honduras 
in  one  extensive  tropical  land.  But  their  separation  from  the 
continent  took  place  at  a  remote  period,  and  they  have  since  been 
broken  up  into  numerous  islands,  which  have  probably  undergone 
much  submergence  in  recent  times.  This  has  led  to  that  poverty 
of  the  higher  forms  of  life,  combined  with  the  remarkable  similarity 
which  now  characterises  them ;  while  their  fauna  still  preserves  a 
sufficient  resemblance  to  that  of  Central  America  to  indicate  its 
origin."  Recently,  the  connection  of  the  West  Indies  with  the 
mainland  has  been  worked  out  more  fully  by  Mr  J.  W.  Spencer2, 
who,  from  observations  made  on  the  buried  river  channels  so 
numerous  in  some  of  the  islands,  concludes  that  there  have  been 
several  epochs  of  connection  with  the  continent,  one  of  which  was 
so  late  in  date  as  the  Plistocene  epoch.  The  extinction  in  the 
islands  of  the  great  majority  of  the  mammals  of  the  continent 
is  attributed  to  drowning. 

1  This  is  not  the  opinion  of  Dr  A.  Agassiz,  who  regards  them  as  oceanic 
islands. 

2  Geological  Magazine,  1894,  pp.  448 — 451. 


III.]  WEST   INDIES.  139 

Here  a  presumed  connection  between  North  and  South 
America  by  way  of  the  West  Indies  must  be  referred  to,  the 
evidence  in  favour  of  which  has  been  summarised  by  Dr  J.  W. 
Gregory1  as  follows: — "It  is  not  at  all  certain  that  when  the 
isthmus  of  Panama  was  submerged  there  was  a  free  communica- 
tion between  the  Atlantic  and  Pacific  Oceans.  The  Caribbean 
Sea  may  then  have  been  a  gulf  from  the  Pacific,  separated  from 
the  Atlantic  by  the  land  area  of  the  hypothetical  'Antillia.'  That 
there  was  once  a  connection  between  North  and  South  America 
along  the  chain  of  the  Windward  Islands,  Cuba,  the  Bahamas,  and 
Florida  is  not  improbable.  Evidence  for  this,  either  in  whole  or 
in  part,  has  been  advanced  by  De  Castro  and  others.  Further 
evidence  could  be  adduced  from  the  study  of  the  land-shells,  and 
also  from  the  remarkable  distribution  of  Peripatus.  That  Cuba 
was  once  connected  with  Yucatan  and  Florida  is  almost  certain ; 
that  this  connection  was  in  existence  in  the  Pliocene,  and  probably 
also  in  the  Plistocene,  is  shown  by  the  evidence  collected  by  De 
Castro.  That  the  area  of  the  Windward  Islands  was  occupied  by 
land  in  the  lower  Tertiary  is  also  most  probable.  But  this  was  all 
submerged  at  the  period  when  the  Oceanic  [Miocene]  deposits  of 
Barbados  were  laid  down.  There  is  no  adequate  evidence  to 
show  that  at  any  time  after  this  was  there  more  land  in  this  region 
than  there  is  at  present."  With  regard  to  the  land-shells,  Mr 
A.  H.  Cooke2  writes  "that  a  certain  number  of  the  characteristic 
North  American  genera  are  found  in  the  Antillean  sub-region, 
indicating  a  former  connection,  more  or  less  intimate,  between  the 
West  Indies  and  the  mainland.... A  small  amount  of  South  Ameri- 
can influence  is  perceptible  throughout  the  Antilles,  chiefly  in  the 
occurrence  of  a  few  species  of  Bulimulus  and  Simpulopsis.  The 
South  American  element  may  have  strayed  into  the  sub-region  by 
three  distinct  routes  :  (i)  by  way  of  Trinidad,  Tobago,  and  the 
islands  northward ;  (2)  by  a  north-westerly  extension  of  Honduras 
towards  Jamaica,  forming  a  series  of  islands,  of  which  the  Rosalind 
and  Pedro  banks  are  perhaps  the  remains;  (3)  by  a  similar 
approximation  of  the  peninsula  of  Yucatan  and  the  western 
extremity  of  Cuba."  This  seems  to  indicate  that  such  Antillean 

1  Quart.  Journ.  Geol.  Soc.  Vol.  LI.  p.  305  (1895). 

2  Cambridge  Natural  History — Mollusca,  pp.  345,  346  (1895). 


140  THE    NEOG^IC   REALM.  [CHAP. 

connection  as  may  have  existed  between  North  and  South 
America  was  of  a  very  incomplete  and  transitory  nature ;  and  that 
before  the  end  of  the  Miocene  there  was  never  any  route  in  this 
direction  by  which  mammals  passed  from  the  one  continent  to  the 
other. 

In  this  connection  it  may  be  mentioned  that  Dr  Hart  Merriam l 
has  proposed  to  unite  Central  America  with  the  West  Indies  to 
form  a  separate  zoological  region — the  Tropical — of  equal  rank 
with  the  Sonoran ;  but  however  much  may  be  urged  in  favour  of 
this  view,  the  multiplication  of  regions  is  much  to  be  deprecated. 

The  practical  or  entire  absence  of  non-volant  native  mammals, 
both  recent  and  fossil,  from  all  the  other  South 

Other  Islands.  .  .....,-  .  . 

American  islands,  with  the  exception  of  1  icrra  del 
Fuego  and  the  Falklands,  properly  excludes  their  consideration 
from  this  volume,  although  it  is  almost  essential  that  a  few  words 
should  be  said  with  regard  to  the  Galapagos  group,  more  especially 
since  conflicting  views  have  been  expressed  concerning  their  rela- 
tions to  the  mainland.  In  respect  to  Tierra  del  Fuego  and  some 
of  the  adjacent  islets,  these  may  really  be  regarded  as  a  part  of 
the  continent,  since  the  main  dividing  channel  is  extremely 
narrow,  and  species  like  the  guanaco  are  common  both  to  the 
islands  and  the  mainland.  And  although  the  Falkland  Islands 
lie  about  350  miles  to  the  eastward  of  southern  Patagonia,  yet 
they  are  separated  by  a  comparatively  shallow  sea  (less  than  a 
hundred  fathoms  in  depth),  and  it  is  thus  evident  that  they  were 
connected  at  no  very  distant  date  with  the  mainland.  Of  the  two 
indigenous  mammals,  the  most  remarkable  is  the  Falkland  Island 
wolf  (Cam's  antarcticus],  which  differs  markedly  from  all  the  Canidce 
of  the  mainland,  and  is  apparently  closely  allied  to  the  North 
American  coyote  (C.  latrans).  The  other  is  a  species  of  groove- 
toothed  mouse  (Rhithrodon).  With  regard  to  Fernando  Noronha, 
the  poverty  of  its  fauna  induces  Mr  Beddard2  to  class  it  among 
oceanic  islands,  although  there  is  some  affinity  between  its  fauna 
and  flora  and  those  of  South  America  and  the  West  Indies. 

Of  far  more  interest  are  the  Galapagos  islands,  situated  on  the 
equator,  at  a  distance  of  about  five  hundred  miles  westward  of  the 

1  Appendix,  No.  19,  p.  33. 

2  Ibid.  No.  5,  pp.  190,  207. 


III.]  GALAPAGOS    ISLANDS.  14! 

coast  of  Ecuador.  Entirely  volcanic  in  structure,  they  are  sur- 
rounded by  a  sea  of  great  depth ;  and  according  to  the  view  both 
of  Wallace  and  Darwin,  they  have  never  been  connected  with  the 
mainland,  while  the  latter  observer  is  also  of  opinion  that  for 
countless  ages  they  have  been  separated  from  one  another.  The 
known  mammals  include  a  bat  (Atalapha],  a  rat  (Afus),  doubtless 
introduced,  and  a  peculiar  species  of  white-footed  mouse  (Sitomys 
bauri).  Of  reptiles  the  islands  contain  two  peculiar  genera  of 
iguanoid  reptiles,  and  no  less  than  five  species  of  giant  tortoises 
belonging  to  the  genus  Testudo.  The  iguanoids  are  nearly  related 
to  South  American  types ;  but  there  are  no  tortoises  now  living 
on  the  mainland,  although  a  large  species  flourished  in  Argentina 
during  the  Pampean  epoch,  while,  as  already  stated,  others  are 
now  found  living  in  the  Mascarene  islands,  and  extinct  species 
occur  in  the  middle  and  later  Tertiary  deposits  of  the  United 
States,  Europe,  and  Northern  India.  It  may  accordingly  be 
taken  for  granted  that  both  the  iguanoid  lizards  and  the  giant 
tortoises  reached  the  island  from  the  South  American  mainland ; 
but  the  question  is  how  did  they  arrive  ?  Dr  Wallace  is  of  opinion 
that  both  were  transported  across  the  sea,  although  by  what  means 
is  unknown.  This  view  is,  however,  disputed  by  Dr  G.  Baur1, 
who  believes  that  the  Galapagos  islands  were  formerly  connected 
not  only  with  one  another,  but  likewise  with  the  mainland.  He 
observes  that  if  the  Galapagos  be  oceanic  islands,  their  inhabitants 
could  only  have  been  introduced  by  accident  from  other  regions ; 
"  but  on  such  a  supposition  we  are  absolutely  unable  to  explain 
the  harmonious  distribution,  we  cannot  explain  why  every,  or 
nearly  every,  island  has  its  peculiar  race  or  species,  not  repre- 
sented on  any  other  island.  If  some  animals  could  be  carried 
over  hundreds  of  miles  to  the  islands,  why  are  they  not  carried 
from  one  island  to  the  other?  But  besides  that,  how  could  we 
make  plain  the  presence  of  such  peculiar  forms  as  the  gigantic 
land-tortoises?  According  to  the  elevation  theory,  we  can  only 
think  of  an  accidental  importation  of  these  tortoises  by  some 
current,  because  they  are  unable  to  swim.  After  the  islands  had 
been  elevated  out  of  the  sea,  it  happened  once,  by  a  peculiar 

1  Proceedings  American  Antiquarian  Society,  Oct.  1891. 


142  THE    NEOG^IC   REALM.  [CHAP. 

accident,  that  a  land-tortoise  was  carried  over.  Alone  it  could 
not  propagate.  This  was  only  possible  after  a  similar  accident 
imported  another  specimen  of  the  same  species,  of  the  other  sex, 
to  the  same  island.  Or  we  could  imagine  that  at  the  same  time 
animals  of  both  sexes  were  thus  accidentally  introduced.  By  this 
we  could  at  least  explain  the  population  of  a  single  island.  But 
how  did  all  the  other  islands  become  populated?  To  explain 
this  we  should  have  to  invoke  a  thousand  accidents.  The  most 
simple  explanation  is  given  by  the  theory  of  subsidence.  All  the 
islands  were  formerly  connected  with  one  another,  forming  a 
single  large  island ;  subsidence  kept  on,  and  the  single  island  was 
divided  up  into  several  islands.  Every  island  developed,  in  the 
course  of  long  ages,  its  peculiar  races,  because  the  conditions  on 
these  different  islands  were  not  absolutely  identical." 

Further  evidence  in  favour  of  the  same  view  is  adduced  by 
Mr  W.  B.  Hemsley1,  who  draws  attention  to  the  marked  simi- 
larity of  the  flora  of  the  Galapagos  Islands  to  that  of  the  South 
American  mainland. 

The  difficulty  of  accounting  for  the  transport  of  reptiles  like 
land-tortoises  across  five  hundred  miles  of  sea  is  undoubtedly  very 
great ;  yet,  in  the  face  of  their  volcanic  nature  and  the  depth  of 
the  surrounding  ocean,  it  is  somewhat  difficult  to  accept  the  view 
that  the  Galapagos  Islands  were  connected  with  South  America. 
In  the  paragraph  quoted  Dr  Baur  appears  to  forget  that  the  first 
tortoise  carried  to  these  islands  may  have  been  a  gravid  female ; 
and  also  that  if  an  ancestral  species  were  established  on  one  of 
the  islands,  there  would  be  nothing  very  wonderful  in  individuals 
being  carried  to  some  of  the  others,  where  they  might  eventually 
differentiate  into  distinct  specific  types.  Moreover,  it  seems  quite 
within  the  bounds  of  possibility  that  the  original  introduction  may 
have  been  effected  by  means  of  eggs  transported  on  natural  rafts. 
That  the  Galapagos  tortoises  were  derived  originally  from  equally 
gigantic  continental  forms,  may  be  taken  for  granted  ;  and  the 
existence  at  the  present  day  of  such  creatures  only  in  these  and 
the  Mascarene  islands  is  one  more  instance  of  the  survival  in  the 
southern  hemisphere  of  ancient  types  which  were  formerly  abun- 

1  Nature,  vol.  lii.  p.  623  (1895). 


III.]  CONCLUSION.  143 

dant  on  the  opposite  side  of  the  equator.  The  practical  absence 
of  mammals  from  the  Galapagos  Islands  is  of  little  import  one 
way  or  the  other,  as  they  might  have  been  drowned  out  during  the 
subsidence;  but  perhaps,  on  the  whole,  a  suspension  of  judgment 
as  to  the  relation  of  these  islands  to  the  mainland  is  the  wisest 
course  to  adopt  at  present. 

Finally,  whether  the  hypotheses  that  have  been  advanced  in 
the  present  chapter  to  explain  the  origin  of  the  peculiar  mammalian 
fauna  of  Neogaea  be  substantiated  or  the  reverse,  there  can  be 
little  doubt  that  Dr  Wallace  has  been  misled  in  his  statement 
that  this  area,  so  "  far  as  we  can  judge  from  the  remarkable 
characteristics  of  its  fauna  and  the  vast  depths  of  the  ocean  east 
and  west  of  it,  has  not  during  Tertiary,  and  probably  not  even 
during  Secondary  times,  been  united  with  any  other  continent, 
except  through  the  intervention  of  North  America." 


CHAPTER    IV. 


THE   ARCTOG^EIC   REALM. 

Features  of  the  Arctogceic  Fauna — Community  of  earliest  Fauna— Evidence  of 
Secondary  Reptiles — Puerco  Fauna — Lemuroids — Insectivora — Carnivores 
— Rodents — Ungulates — Summary  of  the  characteristics  of  the  Mamma- 
lian Fauna  of  Arctogsea. 

ARCTOGSEA,  or  the  Arctogaeic  realm,  includes  the  whole  of  the 
countries  of  the  globe  which  do  not  come  within  the  limits  of 
either  the  Neogaeic  or  Notogseic  realms,  and  thus  embraces  by  far 
the  greater  portion  of  the  land-surface.  Nearly  the  whole  of  this 
vast  tract  lies  to  the  northward  of  the  equator  ;  the  only  portions 
lying  below  that  line  being  the  southern  half  of  Africa,  together 
with  Madagascar  and  some  of  the  Malayan  islands.  As  stated 
in  the  introductory  chapter,  the  term  Arctogaea  was  originally 
proposed  by  Professor  Huxley1,  but,  although  subsequently  used 
in  one  case  by  Dr  Sclater2,  and  at  a  still  later  date  adopted  by 
Dr  Blanford3,  has  failed  to  obtain  general  recognition.  There 
can,  however,  be  no  doubt  that,  so  far  as  mammals  at  least 
are  concerned,  the  whole  of  this  vast  tract  is  entitled  to  hold  only 
the  same  relative  rank  as  each  of  the  realms  treated  of  in  the  two 
preceding  chapters;  and  that  if  we  regard  each  of  the  regions 
into  which  the  area  under  consideration  is  divided  by  Sclater  and 
Wallace  as  equivalent  to  each  of  those  two  realms,  we  have  an 
exceedingly  unequal  series  of  divisions.  Not  only  have  the 
Neogseic  and  Notogseic  realms  no  species  of  mammal  common  to 
one  another,  but  if  we  eliminate  the  genus  Cants,  which  is  of 
comparatively  recent  introduction  into  these  areas,  we  shall  find 

1  Appendix,  No.  18. 

2  Ibid.,  No.  27,  p.  214. 

3  Ibid.,  No.  8,  pp.  76,  77. 


[CHAP,  iv.]  ITS  UNITY.  145 

that  all  the  genera,  and  likewise  most  of  the  families,  together 
with  some  of  the  subordinal  or  even  ordinal  groups  of  mammals 
are  likewise  perfectly  different.  Were  it  not  also  for  the  compara- 
tively recent  union  between  South  and  North  America,  to  which 
allusion  has  been  made  in  the  preceding  chapter,  we  should 
likewise  find  just  as  well  marked  a  distinction  between  the 
mammalian  fauna  of  these  two  countries ;  and,  as  a  matter  of" 
fact,  when  we  go  back  to  the  middle  portion  of  the  Tertiary 
epoch,  we  find  such  a  distinction  actually  existing.  Again,  were 
it  not  for  the  intermediate  connecting  Austro-Malayan  region, 
which  forms,  as  we  have  said,  a  kind  of  zoological  No-man's-land, 
there  would  be  an  equally  stringent  line  of  division  between  the 
Notogasic  realm  and  Asia.  If,  on  the  other  hand,  we  take  the 
different  regions  of  Arctogsea,  we  find  not  only  a  certain  number 
of  species  of  mammals  common  to  two  or  more  regions,  but 
when  we  pass  back  into  the  Tertiary  epoch,  the  whole  faunas  of 
several  of  such  regions  merge  more  or  less  completely  into  one 
another,  instead  of  becoming  more  distinct  than  they  are  at  the 
present  day.  The  lion  and  the  leopard,  for  instance,  are  common 
to  India  and  Ethiopian  Africa,  and  during  the  Plistocene  epoch 
ranged  over  a  considerable  portion  of  Europe ;  while  the  range  of 
the  tiger  includes  not  only  India  and  Ceylon,  but  likewise  a 
considerable  portion  of  Central  Asia  and  China.  The  caracal 
and  the  hunting-leopard  are  also  common  to  India  and  Africa ; 
the  British  fox  ranges  not  only  over  Europe  and  a  large  portion  of 
Asia  north  of  the  Himalaya,  but  likewise  over  a  part  of  North 
America;  and  the  common  otter  is  found  alike  in  India  and 
Europe.  Still  more  numerous  are  the  species  of  mammals  com- 
mon to  Europe,  Northern  Asia,  and  North  America. 

Recapitulating  some  of  the  details  given  in  the  introductory 
chapter,  it  may  be  observed  that  by  Messrs  Sclater  and  Wallace 
the  area  here  included  in  the  Arctogaeic  realm  was  divided  into 
the  Nearctic,  Palaearctic,  and  Oriental  regions.  Professor  Newton, 
who  was  subsequently  followed  by  Dr  Heilprin,  proposed  to 
brigade  the  first  two  of  these  together  under  the  name  of  the 
Holarctic  region.  At  a  still  later  date  Dr  Blanford1,  who  as 

1  Appendix,  No.  8,  p.  76. 
L.  IO 


146  THE   ARCTOG^EIC   REALM.  [CHAP. 

already   stated,    takes   Arctogaea  as    one   of    the    three    primary 
divisions  of  the  globe,  proposed  to  subdivide  it  as  follows,  viz.: — 

1 .  Madagascar. 

2.  Africa,  south  of  the  tropic  of  Cancer. 

3.  Oriental,   South-eastern  Asia,  and  Malayan  islands  to 

Wallace's  line. 

4.  Aquilonian,  Europe,  Asia  north  of  the  Himalaya,  Africa 

north  of  the  tropic  of  Cancer,  and  America  north  of 
about  45°. 

5.  Media-Columbian,  America,  between  about  25°  and  45° 

north  latitude. 

The  importance  of  this  division  was,  firstly,  the  recognition  of 
the  right  of  Madagascar  and  the  adjacent  islands  to  form  a  region  by 
themselves  ;  and,  secondly,  the  separation  of  the  Medio-Columbian 
region  from  the  rest  of  North  America.  And  it  will  be  noted  that, 
if  we  take  away  that  area,  the  Aquilonian  region  corresponds  to 
the  Holarctic  of  Newton  and  Heilprin.  A  further  modification  was 
proposed  in  1892  by  Dr  C.  H.  Merriam1,  who  gave  the  name  of 
Sonoran  region  to  the  area  corresponding  approximately  with  the 
Medio-Columbian  of  Dr  Blanford,  and  suggested  that  the  southern 
portion  of  the  Eastern  Holarctic  region  should  form  a  region  by 
itself;  the  name  of  Boreal  region  being  adopted  for  what  remained 
of  the  Holarctic  after  the  subtraction  of  the  Sonoran  region  and 
a  corresponding  area  in  the  Eastern  Hemisphere. 

Although  from  many  points  of  view  the  retention  of  such 
well-known  terms  as  Palsearctic  and  Nearctic  would  be  a  great 
convenience,  the  close  resemblance  of  the  existing  mammalian 
fauna  of  the  whole  of  northern  Arctogaea  compels  us  to  adopt 
the  view  that  the  area  forms  but  a  single  zoological  region.  For 
this  region  the  name  Holarctic  may  be  retained ;  while  for  the 
southern  portion  of  the  old  Nearctic  region,  the  term  Sonoran  is 
the  most  appropriate.  In  the  Eastern  hemisphere  the  whole  of 
that  portion  of  Arctogsea  not  included  in  either  the  Malagasy, 
Ethiopian,  or  Oriental  regions  is  provisionally  included  in  the 
Holarctic,  although  when  our  knowledge  of  distribution  is  less 

1  Appendix,  No.  19. 


IV.]  FAUNISTIC   CHARACTERS.  147 

imperfect  it  may  subsequently  be  found  practicable  to  separate  a 
distinct  Mediterranean  region. 

In  an  area  of  such  vast  extent  as  the  Arctogseic  realm,  which 
embraces  countries  from  the  equator  to  the  most 
northern  habitable  lands,  it  is,  of  course,  perfectly     the  Arctogseic 
unnecessary   to    say   anything   as   regards    climate 
and   physical  features,  and  we   may  accordingly  proceed   forth- 
with to  discuss  the  leading  features  of  the  mammalian  fauna  as  a 
whole. 

From  the  Notogaeic  realm,  in  its  typical  form,  Arctogaea  is 
distinguished  by  the  absence  of  monotremes  and  diprotodont 
marsupials,  not  only  at  the  present  epoch,  but  so  far  as  we  know, 
in  past  times  also.  From  the  Neogaeic  realm  it  is  equally  well 
differentiated  by  the  absence  at  the  present  day  of  all  the  peculiar 
Neogaeic  types  of  edentates,  and  likewise  at  all  epochs  of  Neo- 
tropical monkeys  and  marmosets.  Such  of  the  former  as  are 
found  in  North  America  are  indeed,  as  we  have  seen  in  the  last 
chapter,  only  intruders  from  the  south  since  the  epoch  of  the 
earlier  Pliocene;  and  in  the  Miocene  the  Arctogaeic  fauna  was 
further  distinguished  from  that  of  Neogaea  by  the  absence  of  the 
peculiar  subordinal  groups  of  ungulates  characteristic  of  the  latter. 
The  Insectivora,  which  with  the  exception  of  the  solenodons  are 
practically  wanting  in  Neogaea,  and  are  unknown  in  Notogaea 
proper,  are  abundant  in  all  the  regions  of  this  realm. 

We  might  almost  go  one  step  further  than  this,  and  say  that 
Arctogaea  previous  to  the  Pliocene  epoch  was  characterised  by 
being  the  sole  habitat  of  almost  all  the  families  of  Eutherian 
terrestrial  mammals,  with  the  exception  of  those  characteristic  of 
the  Santa  Crucian  epoch  of  Neogaea.  But  although  this  would  be 
practically  true,  it  would  land  us  in  the  difficulty  that  the  Ethio- 
pian region  would  probably  have  to  be  excluded  from  Arctogaea, 
seeing  that  the  higher  mammals  of  the  former  region  are  but 
comparatively  recent  immigrants.  Still,  it  may  be  stated  that 
northern  Arctogaea  is  the  original  habitat  of  all  the  modern  types 
of  the  higher  Eutherian  mammals. 

Another  feature  of  Arctogaea  is  the  absence  at  the  present  day 
of  all  marsupials  except  opossums,  while  these  are  only  sparingly 
represented  in  its  western  half.  Moreover,  with  the  exception  of 

10—2 


148  THE   ARCTOG^EIC   REALM.  [CHAP. 


the  same  family  group,  which  are  only  known  from  strata  of  the 
Oligocene  and  Miocene  epochs,  marsupials  appear  to  have  been 
absent  from  a  large  part  of  the  realm  during  the  Tertiary  period, 
although  there  is  reason  to  believe  that  during  the  Eocene  they 
must  have  survived  in  south-eastern  Asia1.  Among  volant 
mammals,  bats  of  the  Neogaeic  family  PhyllostomatidGR*  are  now 
absent  from  the  whole  of  the  realm,  with  the  exception  of  a  part 
of  the  Pacific  side  of  North  America.  Again,  to  revert  to  the 
non-volant  forms,  the  Lemuroid  suborder  of  the  Primates  seems  to 
have  been  absolutely  restricted  to  Arctogaea,  at  least  since  the 
Miocene  epoch,  although  it  may  turn  out  that  the  monkeys  and 
marmosets  of  South  America  may  be  descended  from  ancestral 
lemuroids  which  inhabited  that  country  at  an  epoch  previous 
to  the  deposition  of  the  Santa  Crucian  beds. 

To  take  a  comprehensive  survey  of  the  whole  Secondary  and 

Communit        Tertiary  mammalian  faunas  of  Arctogaea  would  entail 

of  earliest  such    a   mass    of  palaeontological   and   anatomical 

detail  that  it  would  only  weary  the  majority  of  our 

readers,  and  we  must  accordingly  limit  ourselves  to  noticing  some 

of  the  most  striking  features  in  the  earlier  faunas,  and  then  pass 

on  to  the  consideration  of  some  of  the  more  widely  spread  modern 

groups. 

In  the  chapter  devoted  to  the  Notogaeic  realm,  it  has  been 
already  pointed  out  (p.  51)  that  during  the  Jurassic  period  Europe 
and  North  America  were  populated  with  a  fauna  of  polyprotodont 
marsupials  of  small  size,  some  of  which  appear  to  have  been  the 
ancestral  types  from  which  those  now  inhabiting  the  Notogseic 
and  Neogaeic  realms  were  derived,  while  others  have  disappeared 
entirely.  It  will  be  unnecessary  to  recapitulate  the  names  of  the 
more  representative  of  these  forms,  but  it  may  be  stated  that  while 
the  fauna  of  the  lower  Jurassic  Stonesfield  Slate  has  no  equivalent 
in  North  America,  that  of  the  upper  Jurassic  Purbeck  beds  of  Dor- 
setshire is  paralleled  in  the  latter  area.  Although  some  difference 
of  opinion  prevails  among  palaeontologists  as  to  the  identity  of  the 
American  with  the  European  genera,  there  can  be  no  doubt  that 

1  Vide  suprd,  p.  55. 

2  The  genus  Necromantis,  from  the  French  Oligocene,  has  been  assigned 
to  this  family. 


IV.]  COMMUNITY   OF   EARLY   FAUNA.  149 

many  of  them  are  very  closely  allied  indeed,  while  some  are 
probably  inseparable.  Contemporary  with  these  early  marsupials 
were  members  of  the  group  known  as  Multituberculata,  which  are 
probably  more  or  less  closely  related  to  the  existing  monotremes, 
or  egg-laying  mammals,  and  form  with  them  the  subclass  Proto- 
theria.  An  essential  feature  of  these  multituberculates  is  that 
the  molar  teeth  were  divided  by  one  or  more  grooves  into  longi- 
tudinal ridges,  covered  with  numerous  blunt  tubercles ;  such 
grooves  being  very  generally  two  in  number  in  the  upper  molars, 
while  the  lower  teeth  have  but  a  single  one.  Apparently  in  all 
cases  the  extremities  of  the  jaws  were  armed  with  a  pair  of  chisel- 
like  incisor  teeth,  behind  which  in  the  upper  jaw  there  may  have 
been  a  pair  of  smaller  teeth.  Very  generally  the  last  premolar 
tooth,  as  in  the  English  Purbeck  genus  Plagiaulax,  was  com- 
pressed and  trenchant  in  shape,  with  its  upper  edge  regularly 


FIG.  28.     RIGHT  SIDE  OF  LOWER  JAW  OF  Plagiaulax.     Enlarged. 
p.  premolars,  m.  molars. 

convex,  and  its  sides  marked  by  oblique  grooves;  but  in  other 
forms  (Polymastodoti)  this  tooth  was  of  a  more  tubercular  type. 
Without  going  into  disputed  questions,  it  may  be  stated  that  this 
group  was  represented  by  closely  allied  forms  in  the  Jurassic  of 
both  Europe  and  North  America;  while  it  is  also  known,  from 
the  evidence  of  a  single  genus  (Tritylodon)^  to  have  extended  its 
range  to  South  Africa  ;  this  genus  also  occurring  in  the  European 
Trias,  and  thus  affording  another  instance  of  the  wide  range  of  the 
earlier  faunas. 

Although  in  Europe  the  only  known  traces  of  mammalian  life 
during  the  succeeding  Cretaceous  period  occur  in  the  Wealden 
beds  (which  are  the  immediate  successors  of  the  Jurassic  Pur- 
becks),  in  North  America  a  well-developed  fauna  of  polyprotodont 


ISO  THE   ARCTOG^EIC   REALM.  [CHAP. 

marsupials  and  multituberculates  is  met  with  in  rocks  of  Cretaceous 
age ;  and  it  is  most  probable  that  if  suitable  freshwater  beds  were 


FIG.  29.     UPPER  SURFACE  OF  SKULL  OF   Tritylodon.     Somewhat  reduced. 

extant,  the  same  fauna  would  be  found  in  Europe.  By  the 
commencement  of  the  Tertiary  epoch,  most  of  this  old  fauna  seems 
to  have  disappeared;  but  in  the  Puerco  beds  of  the  United  States, 


FIG.  30.     UPPER  MOLAR  OF  A  SMALLER  SPECIES  OF  Tritylodon. 
Natural  size  and  enlarged. 

and  the  equivalent  deposits  of  Cernays,  in  the  south  of  France, 
which  seem  to  form  a  transition  between  the  Secondary  and 
Tertiary,  the  Multituberculata  still  persisted,  and  it  is  noteworthy 
that  one  genus  (Neoplagiaulax]  at  least  was  common  to  the 
northern  half  of  the  eastern  and  western  hemispheres. 

It  thus  appears,  so  far  as  the  available  evidence  permits  of  our 
forming  a  judgment,  that  during  both  the  Jurassic  and  Cretaceous 


IV.]  REPTILIAN    EVIDENCE.  151 

epochs  a  single  mammalian  fauna  was  spread  over  Europe  and 
North  America.  This  being  so,  it  is  a  fair  inference  that  a  similar 
fauna  characterised  a  considerable  portion  of  Asia ;  while  the 
occurrence  of  the  above-mentioned  genus  Tritylodon  points  to  the 
conclusion  that  it  likewise  ranged  over  Africa.  Accordingly,  it 
would  appear  that  not  only  did  the  whole  of  Arctogaea  then  form 
a  single  zoological  realm,  but  that  this  realm  was  indivisible  into 
regions. 

The  evidence  for  this  unity  is,  however,  by  no  means  restricted 
to  mammals,  but  is  supplemented  and  extended  by  £vid 
the  extinct  reptiles  of  the  Secondary  epoch  of  the  Secondary 
earth's  history.  During  the  Triassic  and  early 
Jurassic  periods  there  flourished  an  extinct  ordinal  group  of  rep- 
tiles known  as  the  Anomodontia,  remarkable  for  many  structural 
resemblances  to  mammals,  and  likewise  for  the  peculiarities  of 
their  dentition.  As  a  well-known  example  of  one  section  of  this 
group  may  be  cited  the  dicynodonts,  in  which  the  teeth  were 
reduced,  at  most,  to  a  single  pair  of  tusks  in  the  upper  jaw,  the 
remainder  of  the  jaws  being  ensheathed  in  horn  to  form  a  beak ; 
whereas  Galesaurus  represents  a  second  section  in  which  the  teeth 
simulate  those  of  the  carnivorous  mammals.  These  anomodonts 
are  known  to  have  been  spread  over  Europe,  India,  Africa,  and 
North  America ;  the  dicynodont  types  from  the  three  areas  first 
named  being  so  alike  that  there  is  little  question  that  some  of 
them  were  generically  identical.  The  North  American  forms, 
which  mostly  or  exclusively  come  from  beds  assigned  to  the 
Permian  epoch,  do  not  include  dicynodonts,  but  are  allied  to 
certain  other  African  families,  and  are  also  closely  related  to  their 
European  contemporaries. 

If  we  turn  to  another  order  of  the  same  class,  namely  the 
Dinosauria,  as  represented  by  the  Iguanodon  of  Europe  and  the 
Atlantosaurus  of  the  United  States,  we  find  not  less  well- 
marked  similarities  in  the  Jurassic  and  Cretaceous  fauna  of  the 
whole  of  Arctogaea,  this  group  being  represented  by  closely  allied, 
and  in  many  cases  generically  identical  forms,  not  only  in  Europe, 
India,  and  South  Africa,  but  likewise  in  Madagascar  and  South 
America.  For  instance,  in  that  section  of  the  order  known  as 
the  Sauropoda,  which  includes  the  most  gigantic  forms,  and  is 


152  THE   ARCTOG^EIC   REALM.  [CHAP. 

characterised  by  the  presence  of  large  chambers  in  the  sides  of  the 
vertebrae  of  the  neck  and  trunk,  we  find  not  only  that  several 
genera,  such  as  Morosaurus,  are  common  to  the  upper  Jurassic 
and  lower  Cretaceous  strata  of  Europe  and  the  United  States  ;  but 
we  also  find,  one  genus  (Titanosaurus)  in  India,  Europe,  and 
Patagonia,  while  a  second  (Bothriospondylus)  occurs  in  countries 
as  far  apart  as  England  and  Madagascar1.  Again,  in  the  carni- 
vorous or  Theropodous  section  of  the  order,  as  typified  by  the 
English  Megalosaurns,  we  find  certain  closely  allied  or  identical 
generic  types  common  to  Europe  and  South  Africa.  Further 
evidence  in  the  same  direction  is  afforded  by  the  discovery  in  the 
Jurassic  of  Madagascar  of  a  genus  of  extinct  crocodiles  (Steneo- 
saurus)  which  were  abundantly  represented  during  the  same  epoch 
in  Europe.  Among  the  class  of  fishes  we  have  also  the  genus 
Ceratodus,  now  living  in  Queensland,  represented  in  the  Secondary 
rocks  of  Europe,  India,  Africa,  and  North  America. 

With  regard  to  the  land-fauna  of  Australia  at  the  same  epoch 
we  have  less  evidence ;  anomodonts,  and,  we  believe,  dinosaurs, 
being  unknown  from  that  country.  Among  the  amphibians,  how- 
ever, we  find  in  the  extinct  order  of  Labyrinthodontia  certain 
genera,  such  as  Bothriceps  and  Micropholis,  common  to  Australia 
and  South  Africa,  both  of  these  being  closely  allied  to  the  Indian 
Brachyops. 

This  reptilian  evidence  thus  clearly  points  to  the  conclusion 
that  during  the  greater  part  of  the  Secondary  period  not  only 
had  Arctogaea  a  single  widely-spread  fauna,  but  that  the  same 
fauna  was  represented  in  South  America,  and  at  least  partially  in 
Australia.  Hence  at  this  date  no  zoological  realms  can  be  distin- 
guished, and  it  was  probably  not  till  late  in  Cretaceous  times  that 
Arctogaea  was  differentiated  from  the  rest  of  the  world  as  a  realm. 
Needless  to  say,  the  great  continents  and  islands  during  the 
epochs  in  question  must  have  had  free  communication  with  one 
another,  and  it  is  highly  probable,  as  Dr  Blanford2  suggests,  that 
Madagascar  then  formed  a  line  of  connection  between  Africa  and 
India.  It  is  possible  that  even  at  the  early  part  of  the  Secondary  era, 

1  Possibly  future  discoveries  may  show  differences  worthy  of  generic  distinc- 
tion between  these  forms,  but  this  would  not  affect  the  general  question. 

2  Appendix,  No.  8,  pp.  88,  et  seq. 


IV.]  PUERCO   FAUNA.  153 

"when  South  Africa  was  united  to  India  via  Madagascar  on  one 
side,  and  to  South  America  on  the  other,  especially  if  the  Indo- 
Malay  continent  was  also  connected  with  the  Australian,  there 
may  have  been  a  girdle  of  land,  chiefly  in  low  latitudes,  round 
nearly  three-quarters  of  the  earth's  circumference  from  Peru  to 
New  Zealand  and  the  Fiji  Islands."  The  vertebrate  testimony 
does  not,  however,  countenance  the  idea  that  such  southern  land 
was  cut  off  from  Europe  and  northern  Asia  by  sea.  It  is  true 
that  the  evidence  in  favour  of  such  an  isolation  is  afforded  by  the 
identity  of  the  Carboniferous  (Damuda-Talchir)  floras  of  Australia, 
South  Africa,  Peninsular  India,  and  Central  Argentina1,  and  their 
total  dissimilarity  from  those  of  Europe,  Northern  Asia,  and  North 
America ;  and  it  is  suggested  that  the  same  conditions  may  have 
prevailed  during  the  Jurassic2.  This,  however,  the  vertebrate 
evidence  certainly  does  not  support ;  and  hence,  while  admitting 
the  isolation  of  a  great  southern  (subtropical)  continent  during 
the  Palaeozoic  era,  it  appears  probable  that  since  that  epoch  most 
of  the  southern  lands  have  been  from  time  to  time  more  or  less 
closely  connected  with  those  to  the  north3. 

Leaving  these  difficult  problems  with  the  foregoing  remarks, 
we  pass  on  to  notice  briefly  the  Puerco  mammalian 

Puerco  Fauna. 

fauna  of  the  United  States,  which,  together  with  the 
approximately  equivalent  Cernaysian  fauna  of  Europe,  is  of  especial 
interest  as  showing  a  transition  between  the  Cretaceous  and 
Tertiary.  As  we  have  already  said,  this  fauna  includes  several 
representatives  of  the  multituberculates,  which  are  essentially  a 
Secondary  group,  and  one  of  which  is  common  to  the  Cernaysian 
fauna.  In  addition  to  these,  four  orders  of  eutherian  mammals  are 
represented,  namely  the  Primates,  the  Carnivora,  the  Ungulata, 
and  the  extinct  group  known  as  the  Tillodontia.  It  is,  however, 
very  noteworthy  that  in  all  these  orders  it  is  only  the  lowest 
sections  that  were  in  existence  during  the  Puerco  epoch.  Thus 

1  See  F.  Kurtz,  Rev.  Mus.  La  Plata,  Vol.  vi.  p.  117,  and  Rec.  Geol.  Surv. 
India,  Vol.  xxvni.  p.  in  (1895). 

2  Appendix,  No.  8,  p.  96. 

3  Dr  Blanford  writes  to  me  that  he  believes  the  Palaeozoic  connection  be- 
tween South  America  and  South  Africa  was  tropical  or  subtropical,  rather  than 
antarctic,  and  hence  that  "the  evidence  for  an  antarctic  continent  in  upper 
Mesozoic  or  Tertiary  times  is  very  slight  indeed." 


154  THE   ARCTOGyEIC   REALM.  [CHAP. 


all  the  Primates  belong  to  the  lemuroid  section,  and  include  no 
monkeys  or  apes ;  and  the  carnivores  are  represented  solely  by 
the  extinct  creodont  group,  which  differs  from  the  existing  members 
of  the  order  by  the  simpler  and  more  primitive  structure  of  the 
limbs  and  teeth.  The  ungulates,  again,  belong  exclusively  to  two 
extinct  suborders,  respectively  termed  the  Condylarthra  and  the 
Amblypoda,  both  of  which  are  very  primitive  types,  with  five-toed 
limbs  of  simple  structure,  the  former  still  retaining  evidences  of 
affinity  with  the  early  carnivores.  The  tillodonts  are  quite  unlike 
any  existing  forms,  having  a  pair  of  incisor  teeth  similar  to  those 
of  rodents  in  the  front  of  the  jaws,  while  their  cheek-teeth  recall 
those  of  the  ungulates. 

All  the  Puerco  mammals  are  characterised  by  the  lowness  of 
the  crowns  of  their  molar  teeth,  which  carry  simple  tubercles, 
generally  arranged  in  a  triangle ;  this  type  of  tooth  being  known 
as  the  tritubercular,  and  occurring  in  all  the  orders  found  in  the 
Puerco.  It  will  be  unnecessary  to  mention  the  names  of  the 
genera  occurring  in  this  horizon,  and  it  will  suffice  to  state  that 
while  peculiar  to  these  beds,  many  of  them  belong  to  families  ' 
characteristic  of  the  overlying  Tertiaries.  Thus  we  have  the 
Anaptomorphida  among  the  lemuroids,  the  Arctocyonida,  Mesony- 
chidce,  Proviverridce,  and  Miacida  in  the  carnivores,  and  the 
PhenacodontidcR  among  the  condylarthrous  ungulates.  The  Cernay- 
sian  fauna  is  mostly  represented  by  such  fragmentary  specimens 
that  the  determination  of  the  affinities  of  its  members  is  a  matter 
of  considerable  difficulty;  but  the  forms  were  all  more  or  less 
nearly  allied  to  those  of  the  Puerco,  and  the  creodont  genus 
Dissacus  is  common  to  the  two  formations ;  while  Arctocyon, 
which  is  met  with  in  the  Cernaysian,  also  occurs  in  higher  horizons 
both  in  Europe  and  America. 

A  very  remarkable  fact  connected  with  the  Puerco  fauna  is 
that  out  of  the  39  generic  types  by  which  it  is  represented,  only 
eight  are  followed  by  analogous  forms  in  the  overlying  Wahsatch 
beds,  three  of  which  became  extinct  in  the  still  higher  Bridger 
deposits.  This  leads  Messrs  Osborn  and  Earle  to  the  conclusion 
that  this  early  mammalian  fauna  was  a  kind  of  failure  as  regards 
development,  and  that  only  a  few  of  its  less  specialised  members 
persisted  to  give  rise  to  the  mammals  of  later  periods. 


IV.]  LEMUROIDS.  155 

With  the  Puerco  and  Cernaysian  faunas  we  take  leave  of  the 
Secondary  multituberculates,  and  as  we  ascend  in 

.  .  ...  .  Lemuroids. 

the  Tertiary  series  we  find  a  gradual  and  progressive 
modification  of  the  eutherian  mammals  towards  the  modern  types. 
In  the  ungulates  especially  the  modification  displays  itself  in  the 
more  complex  structure  of  the  molar  teeth,  and  in  the  reduction  of 
the  number  of  toes ;  the  culmination  of  the  latter  line  of  develop- 
ment being  reached  in  the  modern  horses  among  the  perissodactyle 
section  of  the  order,  and  in  the  ruminants  among  the  artiodactyles. 
As  regards  the  molar  teeth,  the  chief  features  are  a  lengthening  of 
the  crowns  in  the  more  specialised  later  forms,  accompanied  by 
complex  infoldings  of  the  surface  of  the  crown  and  sides,  whereby 
the  short-crowned,  or  brachydont  type,  as  exemplified  in  the  tapirs, 
has  developed  into  the  tall,  or  hypsodont  type  characteristic  of  the 
horses.  Instead,  however,  of  tracing  the  succession  of  the  various 
faunas,  it  will  suit  our  present  purpose  better  to  refer  to  the 
distribution  of  the  more  widely  spread  groups  which  are  either 
characteristic  of  Arctogsea  as  a  whole,  or  which  were  common  to 
that  realm  together  with  Neogsea  during  the  Plistocene  period. 

The  lemuroids,  which  are  at  present  unknown  beyond  the 
limits  of  this  realm,  are  first  met  with  in  the  Puerco  beds,  where 
they  are  represented  by  Indrodon,  and  it  is  probable  that  the 
Cernaysian  mammal  described  as  Plesiadapis  (of  which  the  upper 
cheek-teeth  are  shown  in  the  annexed  figure)  belongs  to  the  same 


FlG.    31.      THE    RIGHT    UPPER    CHEEK-TEETH    OF    Plesiadapis : 

p.  premolars,  m.  molars. 

group.  It  will  be  observed  that  in  the  latter  genus  the  molars  are 
of  the  tritubercular  type;  the  same  being  the  case  in  Anapto- 
morphus  of  the  lower  or  Wahsatch  Eocene  of  America.  In  other 
forms,  however,  as  in  Hyopsodus  and  Pelycodus  of  the  lower  Eocene 
of  North  America,  and  probably  also  of  the  European  Eocene,  as 
well  as  in  Microchcerus  of  the  Oligocene  of  France  and  England, 
the  upper  molar  teeth  have  squared  crowns,  and  thus  approximate 


156  THE   ARCTOG^IC   REALM.  [CHAP. 

to  those  of  modern  lemurs.  These  early  forms  differ  however  from 
the  latter  in  that  the  first  of  the  three  lower  premolar  teeth  does 
not  assume  the  form  and  functions  of  a  canine.  Another  well- 


FlG.  32.       RIGHT  UPPER  CHEEK-TEETH  OF  TWO  SPECIES  OF  THE]  LEMUROID 

GENUS  Microchtxrus.     Nat.  size  and  enlarged. 

known  European  lemuroid  is  Adapts,  of  the  European  Oligocene, 
differing  from  all  living  forms  in  having  four  pairs  of  premolar 
teeth. 

With  the  Oligocene,  lemuroids  seem  to  have  disappeared  from 
western  Europe,  and  they  apparently  ceased  to  exist  about  the 
same  date  in  North  America,  after  which  the  entire  order  of  the 
Primates  is  unrepresented  in  the  latter  country.  At  the  present 
day,  as  we  shall  see,  lemuroids  are  confined  to  the  Malagasy, 
Ethiopian,  and  Oriental  regions ;  but  at  what  epoch  the  southern 
migration  took  place  cannot  yet  be  determined. 

Omitting  mention  of  the  bats,  we  pass  on  to  the  Insectivora, 
among  which  we  have  the  mole  family  (Talpidce) 
distributed  over  the  whole  of  the  Holarctic  as  well  as 
the  Sonoran  region,  although  all  the  genera  but  one  are  distinct 
on  the  two  sides  of  the  Atlantic ;  the  single  common  type  being 
the  shrew-moles  (Urotrichus\  which  have  one  species  in  Japan 
and  another  in  the  United  States,  thus  affording  an  instance  of  the 
near  affinity  of  the  fauna  of  eastern  Asia  to  that  of  North  America. 
The  earliest  known  fossil  forms  which  have  been  assigned  to  the 
typical  genus  Talpa  occur  in  the  upper  Oligocene  strata  of  Europe, 
while  in  the  middle  Oligocene  the  family  is  represented  by  the 
allied  Amphidozotherium  (Protalpa).  The  shrews  (Soriddce)^ 
which  likewise  date  from  the  Oligocene  of  the  Continent,  range 
over  the  whole  realm,  and  also  enter  the  Austro-Malayan  region 


IV.]  CARNIVORA.  157 

as  well  as  the  Mexican  subregion,  although  they  are  represented 
in  Madagascar  only  by  a  single  species  of  the  widely  spread  genus 
Crocidura.  Unknown  in  America,  the  latter  genus,  which  includes 
the  well-known  musk-shrews,  is  widely  spread  over  Europe,  Asia, 
and  Africa,  extending  as  far  east  as  Amurland ;  but  the  typical 
genus  Sorex  is  practically  confined  to  the  Holarctic  region1,  while 
other  genera  have  a  more  local  distribution. 

With  the  exception  of  the  civet  tribe  ( Viverridce]  and  hyaenas 
(Hyanidce),  which  are  unknown  in  the  New  World, 

.......  .  Carnivora. 

the  majority  of  the  existing  families  of  the  Carnivora 
have,  if  we  except  Notogaea,  a  cosmopolitan  distribution,  while  in 
many  cases  this  extensive  distribution  also  holds  good  with  regard 
to  genera.  In  Europe  the  Felidce  and  Canidce,  together  with  the 
Mustelida  seem  to  have  made  their  first  appearance  in  the  lower 
Oligocene,  when  they  were  accompanied  by  the  extinct  creodonts ; 
while  in  America  the  two  former  are  known  from  the  John  Day 
group,  corresponding  to  the  European  Miocene.  The  bears 
( Ursidce)  are,  however,  a  later  group,  being  unknown  before  the 
Pliocene.  Although  the  whole  of  the  families  mentioned  above 
are  represented  in  South  America  at  the  present  day  and  during 
the  Plistocene,  they  are,  as  we  have  seen  in  the  last  chapter, 
unknown  in  the  presumably  Miocene  Tertiaries  of  Patagonia,  and 
they  are  therefore  originally  of  Arctogaeic  origin.  Although  most 
of  the  extinct  American  Tertiary  genera  of  cats  are  distinct  from 
those  of  Europe,  the  sabre-toothed  tigers  (Machcerodus)  were 
common  to  both  areas,  and  likewise  ranged  into  Notogaea;  and 
the  existing  Felis  has  a  similar  cosmopolitan  distribution.  A  more 
.specialised  sabre-toothed  genus  (Eusmilus)  is  likewise  common 
to  North  America  and  Europe.  As  examples  of  extinct  American 
cats,  we  may  name  Nimravus  and  Archcelurus ;  while  the  Oligocene 
^Elurictis  may  be  cited  as  an  Old  World  form.  The  aforesaid 
distinction  between  the  Oligocene  and  Miocene  Felidcz  of  North 
America  and  Europe  is,  however,  an  indication  that  by  this  date 
the  mammalian  faunas  of  Western  and  Eastern  Arctogaea  had 
become  differentiated  to  a  certain  extent,  although,  as  now,  there 
were  many  types  common  to  the  two  areas. 

1  It  has  one  species  in  the  Sonoran. 


158  THE   ARCTOG^IC   REALM.  [CHAP. 

Much  the  same  story  is  told  by  the  fossil  dogs  ( Canidce)  of  the 
two  areas.  In  the  Miocene  of  North  America  we  meet  with  the 
genus  Temnocyon,  characterised  by  the  cutting  heel  of  the  lower 
carnassial  tooth;  while  the  more  civet-like  Cynodictis  appears  to  be 
confined  to  the  Tertiaries  of  Europe.  The  bear-like  genus  Am- 
phicyon,  differing  from  modern  dogs  by  its  plantigrade  feet,  is 
confined  to  the  European  Oligocene  and  Miocene  and  lower 
Pliocene,  but  is  represented  in  the  Miocene  of  America  by  the 
nearly  allied  Daphcemis.  Through  the  intervention  of  the  still 
larger  Dinocyon  of  the  European  Miocene,  the  foregoing  groups  are 
intimately  connected  with  the  bears  (Ursidcz)  by  means  of  the 
genus  Hycenarctus,  which  is  common  to  the  Miocene  and  Pliocene 
of  Europe  and  the  Pliocene  of  India;  and  this  suggests  that  the 
bears  are  originally  an  Old  World  group,  which  have  subsequently 
migrated  into  America.  As  to  whether  true  dogs  (Cam's)  and  cats 
(Felis)  originated  in  America  or  Europe,  we  have  no  means  of 
deciding1.  The  large  weasel  family  (Mustelidce)  calls  for  no  special 
mention  here,  although  its  comparative  poverty  in  South  America 
proclaims  its  Arctogaeic  origin. 

The  community  between  the  mammalian  faunas  of  Eastern 
and  Western  Arctogaea  is  perhaps  better  exemplified  by  the  ex- 
tinct creodont  carnivores,  since  none  of  the  families  occurring  there 
have  been  definitely  recognised  in  the  South  American  area. 
Differing  from  modern  carnivores  by  the  absence  of  a  pair  of 
differentiated  carnassial  teeth  in  each  jaw,  as  well  as  by  the 
scaphoid  and  lunar  bones  of  the  wrist  generally  remaining 
separate,  and  by  the  nearly  flat  upper  surfaces  of  the  astragalus  in 
the  ankle,  these  creodonts  make  their  first  appearance  in  the 
Puerco,  and  mostly  died  out  in  the  Oligocene,  although  a  few  seem 
to  have  survived  till  the  Miocene.  In  the  typical  family  Hyczno- 
dontida  we  find  the  genus  Hycenodon  common  to  the  Oligocene  of 
both  sides  of  the  Atlantic,  while  the  European  Pterodon  seems  to 
have  a  transatlantic  representative  in  the  so-called  Hemipsalodon  of 

1  Scott,  Trans.  Amer.  Phil.  Soc.  xvn.  p.  75,  concludes  that  the  evolution 
of  Cam's  took  place  in  North  America,  the  ancestry  being  traced  through 
Cynodesmtts  of  the  John  Day  Beds  to  Daphcemis  of  the  White  River,  and 
thence  to  the  creodont  Miacis  of  the  Bridger ;  Cynodictis  forming  a  lateral 
branch. 


IV.]  RODENTS.  159 

Canada.  Oxycena,  again,  is  found  both  in  America  and  Europe, 
although  in  a  lower  horizon  in  the  former  than  in  the  latter.  In  a 
second  family  (ProviverridcR)  the  typical  genus  Proviverra  is  met 


m 

FlG.    33.      RIGHT   UPPER  MOLARS   OF   ArctoCVOH. 

with  in  the  Bridger  Eocene  of  America,  and  the  French  Oligocene; 
while  in  the  Arctocyonidce,  in  which  the  upper  molar  teeth  are 
bluntly  tritubercular,  Arctocyon  of  the  lowest  Eocene  of  Europe  is 
represented  by  two  allied  genera  in  the  American  Puerco,  one  of 
which  has  been  described  as  Clcenodon. 

In  the  rodents  there  are  three  more  or  less  widely  distributed 
existing  families  restricted  to  Arctogaea.  Of  these, 
the  jerboas  and  their  allies  (DipodidtzY  occur  in  all 
the  regions  with  the  exception  of  the  Malagasy,  Oriental,  and 
Sonoran, — although  the  genera  from  the  different  areas  are  more 
or  less  markedly  distinct.  The  other  two,  namely  the  picas  or 
tailless  hares  (Lagomyida)  and  the  beavers  (Castoridce)  are  now 
severally  represented  by  a  single  genus  confined  to  the  Holarctic 
region.  The  picas  date  from  the  Oligocene  of  Europe,  and  the 
family  not  improbably  originated  in  eastern  Arctogaea ;  while  the 
beavers  have  fossil  representatives  in  both  hemispheres,  with  the 
Miocene  and  Pliocene  genus  Chalicomys  common  to  the  two. 

Although  members  of  the  typical  genus  range  into  the  Neogaeic 
realm  as  far  south  as  Paraguay,  the  squirrel  family  (Sciuridce)  may 
be  regarded  as  a  typical  Arctogaeic  one,  the  ground-squirrels 
(Tamias),  marmots  (Arctomys)  and  susliks  (Spermophilus}  being 
restricted  to  the  Holarctic  region,  though  others  range  over  the 

1  It  has  been  suggested  by  Dobson  that  the  Dipodidcz  are  Hystricomorpha. 
This,  however,  is  disproved  by  Dr  Scott  (P.  Ac.  Philad.  1895,  pp.  269 — 286), 
who  finds  in  the  Uinta  Oligocene  genus  Protoptychus  an  ancestral  type  of  the 
family,  which  thus  appears  to  be  of  N.  American  origin.  From  this  family  are 
probably  descended  the  Geomyidce. 


l6o  THE   ARCTOG^EIC    REALM.  [CHAP. 


whole  of  the  tropical  and  temperate  parts  of  the  realm  with  the 
exception  of  Madagascar.  Remains  of  Sperm  ophilus  and  Sciurus 
are  met  with  in  the  later  Tertiaries  of  Europe';  and  the  extinct 
Plesiarctomys,  which  is  common  to  the  Oligocene  and  Miocene  of 
Europe  and  North  America,  seems  to  be  a  connecting  form 
between  the  squirrels  and  marmots,  having  upper  molar  teeth  of 
the  tritubercular  type. 

As  regards  the  cosmopolitan  family  Muridcz,  including  the 
rats,  voles,  lemmings,  etc.,  it  will  suffice  to  say  that  originally  it 
was  undoubtedly  Arctogaeic ;  the  forms  respectively  inhabiting  the 
Neogaeic  and  Notogaeic  realms  being  comparatively  recent  immi- 
grants. Both  the  subfamilies  of  the  voles  (Microtince)  and  the 
CricetincB  are  common  to  the  entire  Holarctic  region ;  the  latter 
being  represented  in  the  eastern  half  by  the  hamsters  (Cricetus) 
and  in  the  western  by  the  white-footed  mice  (Sitomys],  while  they 
are  the  sole  rodents  inhabiting  Madagascar,  and  have  one  species 
in  Ethiopian  Africa,  where  there  is  also  the  closely  allied  Deomys, 
forming  a  subfamily  by  itself.  The  cricetines  are  indeed  evidently  a 
primitive  type,  which  in  the  Old  World  have  been  largely  supplanted 
or  driven  south  by  the  more  specialised. Murince  (true  rats  and  mice) ; 
but  as  these  are  represented  in  the  Middle  Tertiaries  of  both 
eastern  and  western  Arctogaea,  it  is  difficult  to  decide  which  was 
their  original  habitat.  Little  need  be  said  in  regard  to  the  hares 
(Leporidce),  except  that  they  range  over  the  whole  of  Arctogasa, 
and  have  two  outlying  representatives  in  Neogaea,  which  are 
doubtless  comparatively  recent  immigrants,  although  one  is  known 
to  have  inhabited  Brazil  since  the  Plistocene. 

A  not  less  marked  feature  of  Arctogaea  is  the  absence  of  most 
of  the  Neogaeic  rodent  families  noticed  in  the  preceding  chapter. 
The  existence  of  members  of  one  of  these  (Octodontida)  in  Africa 
is  mentioned  in  the  same  place1,  where  a  reference  to  the  occur- 
rence of  allied  forms  (Theridomyidce]  in  the  European  Tertiaries 
will  also  be  found2. 

One  of  the  most  important  features  in  connection  with  the 
Arctogaeic   ungulates  is    the   total  absence    of  the 

Ungulates.  5  6 

peculiar    subordmal    groups    characteristic   of    the 
1  See  also  the  chapter  on  the  Ethiopian  region.         2  Supra,  p.  86. 


IV.]  ARTIODACTYLES.  l6l 

South  American  Tertiaries ;  although,  as  stated  in  the  last  chapter, 
there  are  indications  of  a  distant  affinity  between  these  and  some 
of  the  primitive  early  European  perissodactyles.  So  far  as  our 
present  knowledge  goes,  both  the  perissodactyle  and  artiodactyle 
suborders  made  their  appearance  in  North  America  during  the 
lower  or  Wahsatch  Eocene  ;  the  former  group  at  least  also  dating 
from  the  same  epoch  in  Europe,  where  the  genus  Hyracotherium, 
which  is  common  to  the  Bridger  and  Wahsatch  Eocene,  and  is  one 
of  the  earliest  ancestors  of  the  horses,  occurs  in  the  London  Clay. 
Commencing  with  the  Artiodactyla,  or  even-toed  group — 
characterised  by  the  toes  corresponding  to  the  third  and  fourth 
of  the  typical  pentedactyle  limb  being  symmetrical  to  a  line  drawn 
between  them — and  taking  into  consideration  only  such  families  as 
have  a  wide  distribution,  we  have  first  to  do  with  certain  extinct 
types  which  serve  to  connect  the  pigs  with  the  ruminants.  The 
most  pig-like  of  these  are  the  animals  forming  the  family  Chcero- 
potamidce,  characterised  by  their  broad  upper  molar  teeth  carrying 
five  blunt  tubercles,  three  of  which  are  on  the  front  half  of  the 
crown.  Although  the  typical  genus  Chosropotamus  appears  to  have 
been  confined  to  the  lower  Oligocene  of  Europe,  the  much  larger 
animals  known  as  Elotherium  (fig.  35)  were  common  to  the  upper 
Oligocene  of  both  hemispheres.  Nearly  allied  is  the  family  of  the 
Anthracotheriidce,  in  which  the  low  tubercles  of  the  molars  assume 


FlG.    34.      RIGHT    UPPER   MOLAR   OF  AncoduS. 

a  more  or  less  crescentic,  or  selenodont  structure,  thus  foreshadow- 
ing those  of  the  ruminants.     Here,  again,  the  typical  genus  is 
restricted  to  the  Old  World,  but  the  nearly  allied  Ancodus  (Hyo- 
L  II 


162 


THE   ARCTOG^IC   REALM. 


-C 


IV.]  ARTIODACTYLES.  163 

potamus]  has  the  same  approximate  distribution  as  Elotherium, 
although  it  also  occurs  in  the  Miocene  of  northern  India,  together 
with  Anthracotherium  and  a  genus  (Tetraconodon]  closely  allied  to 
Elotherium. 

A  group  of  smaller  Eocene  and  Oligocene  mammals,  con- 
stituting the  family  Ccenotheriidce,  differ  from  the  preceding  in 
that  their  upper  molars  have  two  cusps  on  the  front,  and  three  on 
the  hinder  half  of  the  crown.  Here  none  of  the  genera  are  com- 
mon to  the  two  sides  of  the  Atlantic,  Ccenotherium  and  Dichobumis 
being  European,  while  the  latter  is  represented  in  the  Bridger 
Eocene  by  the  closely  allied  Homacodon.  It  has  been  suggested 
that  this  group  includes  the  ancestors  of  the  camel  tribe. 

The  latter  group  (Camelidce),  now  represented  only  by  the 
camels  (Camelus)  in  the  Old  World,  and  the  guanacos,  vicunas, 
and  their  domesticated  allies  the  llamas  in  South  America,  was 
formerly  widely  distributed  in  Arctogaea,  which  was  doubtless  its 
original  home,  since,  as  we  have  seen  in  the  preceding  chapter, 
the  llamas  and  their  allies  are  but  comparatively  recent  immigrants 
into  Neogaea.  Camelus  itself  is  represented  in  a  fossil  state  in  the 
Pliocene  of  northern  India  and  the  Plistocene  of  Algeria ;  but  no 
other  Old  World  representatives  of  the  family  are  known.  Very 
different,  however,  is  the  case  with  North  America,  where,  from 
the  Plistocene  downwards,  we  meet  with  a  host  of  extinct  types, 
such  as  Pliauchenia,  Procamelus,  Protolabis,  etc.,  gradually  con- 
necting the  existing  forms  with  a  small  animal  from  the  middle 
Oligocene  known  as  Poebrotherium,  which  exhibits  many  very 
generalised  characters.  A  still  earlier  representative  of  the  family 
in  Leptotragulus,  of  the  lower  or  Uinta  Oligocene,  which  itself  may 
be  sprung  from  the  aforesaid  Homacodon  of  the  underlying  Bridger 
beds.  It  is  thus  perfectly  evident  that  the  cameloids  were  ori- 
ginally a  N.  American  group.  One  branch  crossed  the  area  now 
occupied  by  Bering  Strait  to  found  the  camels  of  the  Old  World ; 
while,  probably  at  a  later  date,  a  second  branch  passed  over  the 
isthmus  of  Panama  to  persist  in  the  guanacos,  vicunas,  and 
llamas  of  South  America.  The  disappearance  of  the  whole  group 
from  the  northern  half  of  the  New  World  is  a  very  remarkable  fact, 
but  is  paralleled  by  that  of  the  elephants,  rhinoceroses,  lemurs,  and 
several  other  groups. 

II — 2 


164  THE   ARCTOG^EIC    REALM.  [CHAP. 

The  Tragulidce,  or  chevrotains,  which  form  a  group  distinct 
both  from  the  cameloids  and  the  true  ruminants,  are  now  confined 
to  the  Oriental  and  Ethiopian  regions,  being  represented  in  the 
former  area  by  the  true  chevrotains  (Tragulus),  and  in  the  latter 
by  the  single  existing  species  of  water-chevrotain  (Dorcatheriuni). 
Representatives  of  the  latter  genus  occur,  however,  in  the  Miocene 
and  Pliocene  strata  of  Europe ;  while  in  the  Oligocene  we  meet 
with  the  more  generalised  extinct  genera  Prodremotherium  and 
Bachitheriuwi.  In  North  America  the  group  is  but  poorly  repre- 
sented, and  apparently  confined  to  the  White  River  Oligocene, 
where  we  find  two  types  described  under  the  names  of  Leptomeryx 
and  Hypertragulus ;  the  latter  differing  from  the  existing  forms  by 
having  both  the  third  and  fourth  metacarpals  and  the  correspond- 
ing metatarsals  separate,  instead  of  being  fused  together  to  form  a 
cannon-bone.  It  is  difficult  to  decide  whether  the  group  was 
originally  an  Old  or  a  New  World  one ;  but,  on  the  whole,  it  is 
probable  that  it  originated  in  the  former  area. 

We  now  come  to  the  true  ruminants,  or  Pecora,  forming  the 
most  specialised  group  of  all  the  artiodactyle  section  of  the  ungu- 
lates, and  characterised  by  the  completely  crescentic,  or  selenodont 
conformation  of  the  columns  of  their  molar  teeth,  which  are 
frequently  of  great  height ;  and  likewise  by  the  fusion  of  the  third 
and  fourth  metacarpals  and  metatarsals  into  a  cannon-bone,  and 
by  the  imperfect  development  or  disappearance  of  the  lateral 
metacarpals  and  metatarsals.  In  the  family  of  the  Cervida  the 
typical  deer,  or  those  included  in  the  genus  Cervus,  are  almost 
exclusively  Arctogseic1,  being  found  in  all  the  regions  of  the  realm, 
except  the  Sonoran,  Ethiopian,  and  Malagasy.  The  reindeer 
(Rangifer]  and  elk  (Alces]  are  also  solely  Arctogseic  forms,  but 
have  a  more  restricted  range,  being  confined  to  the  more  northern 
portions  of  the  Holarctic  region.  A  more  striking  case  is  afforded 
by  the  hollow-horned  ruminants,  or  Bovidce,  the  whole  of  the 
numerous  members  of  which  are  confined  to  the  realm  under  con- 
sideration, with  the  sole  exception  of  the  anoa  (Bos  depressicornis] 
of  Celebes ;  and  even  the  latter,  as  we  have  seen  in  an  earlier 
chapter,  is  very  closely  related  to  certain  extinct  Indian  forms.  It 

1  The  only  exception  is  Cervus  timoriensis,  which  may  have  been  introduced 
into  its  present  habitat. 


IV.]  PERISSODACTYLES.  165 

may  be  noticed,  however,  that  Bovidce  are  much  more  numerously 
represented  in  Eastern  than  in  Western  Arctogaea  ;  the  latter  area 
having  only  the  genera  Bos,  Ovibos,  Ovis,  and  Haploceros,  each 
with  a  single  species  confined  to  the  more  northern  portions  of 
the  continent ;  the  last  genus  being  peculiar  to  this  area. 

We  now  pass  to  the  perissodactyle,  or  odd-toed  ungulates, 
which  while  agreeing  with  the  artiodactyle  section  in  the  inter- 
locking arrangement  of  the  bones  of  the  wrist  and  ankle  joints, 
and  the  pulley-like  upper  surface  of  the  astragalus  bone  in  the 
latter,  differ  by  the  third  or  middle  toe  and  its  supporting  meta- 
carpal  or  metatarsal  bone  being  symmetrical  in  itself,  and  larger 
than  the  lateral  ones,  when  such  are  retained.  In  this  suborder 
the  family  of  the  tapirs  (Tapiridce),  although  now  mainly  Neogaeic, 
with  one  outlying  Malayan  species,  was  formerly  widely  spread  in 
northern  Arctogaea,  fossil  remains  belonging  to  the  single  existing 
genus  Tapirus  being  abundant  in  the  Pliocene  of  Europe,  although 
none  appear  to  have  been  recorded  from  North  America.  In 
both  Europe  and  America  there  occurs,  however,  the  ancestral 
genus  Protapirus ;  its  remains  having  been  obtained  in  the  former 
area  from  the  upper  Oligocene  phosphorites  of  France,  and  in  the 
latter  from  the  nearly  equivalent  Uinta  beds.  Possibly  a  doubtful 
form  {Palceotapirus)  from  the  middle  Eocene  of  France  should 
also  be  included  in  the  same  family.  The  Uinta  and  Bridger 
deposits  have  also  yielded  a  more  or  less  nearly  allied  form  known 
as  Isectolophus,  which  apparently  also  occurs  in  the  European 
Eocene,  where  it  has  been  described  as  Lophiodon.  Indeed,  in  our 
view,  both  this  genus,  and  the  still  earlier  American  Systemodon, 
which  appears  to  have  been  the  earliest  known  representative  of 
the  tapiroid  stock,  may  be  included  in  the  family  Lophiodontidce, 
where  we  should  also  place  the  ancestral  types  of  the  horses.  In 
this  family  the  genus  Lophiodon,  as  now  restricted,  seems  to  have 
been  confined  to  the  Eocene  of  Europe,  where  it  died  out  without 
giving  rise  to  descendants,  the  same  being  the  case  with  the 
allied  Eocene  genus  Helaletes^.  The  well-known  Hyracotherium, 
which  was  an  animal  of  the  size  of  a  fox  first  described  from 
the  London  Clay  but  subsequently  recorded  from  the  North 

1  Recorded  from    Europe   by  Osborn   and  Wortman,  Bull.  Amer.  Mus. 
Vol.  vii.  p.  360  (1895). 


1 66  THE   ARCTOG^IC   REALM.  [CHAP. 

American  Eocene,  is,  however,  the  proximate  ancestor  of  the 
horse-family  (Equid<z)\  and  we  have  thus  evidence  that  the  fore- 
runners of  both  the  horses  and  tapirs  were  widely  spread  over  the 
whole  of  northern  Arctogaea.  Hyracotherium,  it  may  be  observed, 
had  the  typical  forty-four  teeth  characteristic  of  the  earlier  euther- 
ians,  and  four  toes  to  the  front,  and  three  to  the  hind  feet;  but  in 
the  still  earlier  Phenacodus,  which  seems  to  be  the  ultimate 
ancestor  of  the  horses,  each  foot  was  furnished  with  five  complete 
toes.  As  other  instances  of  the  community  between  the  early 
Tertiary  mammalian  fauna  of  the  northern  halves  of  the  two 
hemispheres,  we  may  cite  the  genus  Pachynolophus  of  the  middle 
and  upper  Eocene  of  Europe  and  the  Bridger  Eocene  of  the  United 
States,  which  connects  Hyracotherium  with  the  under-mentioned 
horse-like  animals ;  and  also  Hyrachyus,  typically  from  the 
Bridger,  but  probably  occurring  in  the  French  phosphorites ;  the 
latter  being  more  nearly  related  to  the  rhinoceroses. 


FlG.    36.      LEFT    UPPER    CHEEK-TEETH    OF    Palceotheri 


Passing  by  several  less  important  genera  confined  to  one  or 
the  other  hemispheres,  we  come  to  the  family  Pal&otheriidce, 
which  is  an  ill-defined  one  including  forms  connecting  the  pre- 
ceding with  the  undoubted  Equidce.  Here  the  typical  Oligocene 
genus  Pal&otheriuni)  which  includes  large  tapir-like  animals  with 
three  toes  to  each  foot,  is  exclusively  European.  The  same  is  the 
case  with  the  contemporary  Anchilophus,  represented  by  smaller 
forms  with  more  decidedly  horse-like  affinities;  but  with  the 
Miocene  and  upper  Oligocene  Anchitherium,  used  in  its  wider 
sense,  we  once  more  come  to  a  genus  common  to  Western  and 
Eastern  Arctogaea.  In  this  genus  the  jaws  are  provided  with  the 
typical  forty-four  teeth;  but  the  last  lower  molar  has  generally  lost 
the  third  lobe  found  in  the  preceding  forms ;  the  fifth  metacarpal 
bone  being  still  represented  by  a  splint.  In  the  small  A. 


IV.] 


PERISSODACTYLES. 


I67 


(Mesohippus)  bairdi  of  the  White  River  Oligocene  of  the  United 
States,  the  incisor  teeth  lack  the  infoldings  of  the  summit  of  the 
crown  characteristic  of  the  horses,  and  the  lateral  digits  are 
relatively  large ;  the  whole  size  of  the  creature  being  comparable 


FIG.  37.     LEFT  UPPER  CHEEK-TEETH  OF  Anchitherium. 

to  that  of  a  Newfoundland  dog.  On  the  other  hand,  in  the  typical 
A.  aurelianense,  from  the  European  Miocene,  the  summits  of  the 
incisors  were  infolded,  as  in  the  horses.  In  spite  of  this  resem- 
blance, Professor  Scott,  from  the  structure  of  the  limbs,  is  of 
opinion  that  the  latter  species  was  not  an  ancestor  of  the  modern 
horses,  but  that  this  position  was  occupied  by  A.  bairdi. 

Restricting  the  term  Equidce  to  those  members  of  the  suborder 
in  which  the  crowns  of  the  cheek-teeth  are  very  tall  (Jiypsodonf), 
with  complicated  infoldings  of  their  enamel,  and  the  hollows  thus 
formed  completely  filled  with  cement,  we  have  in  the  lower 
Pliocene  of  North  America  the  three-toed  genus  Protohippus, 


FlG.    38.       RIGHT   UPPER    MOLAR   TOOTH   OF   A   HORSE    (Equus). 


distinguished  from  the  modern  horses  by  the  shorter  crowns  of  the 
cheek-teeth.  The  widely-spread  genus  Hipparion  differs  in  having 
the  anterior  inner  column  of  the  upper  molars  completely 


1 68 


THE   ARCTOG^IC   REALM. 


[CHAP. 


detached1;  the  feet  being  generally  three-toed,  although  in  one 
Indian  species  the  lateral  digits  are  wanting.  These  three-toed 
horses  are  met  with  in  the  Pliocene  of  Europe,  Asia,  and  North 


FlG.    39.       UNDER    SURFACE    OF    SKULL    OF   Hipparion. 

America  ;  and  it  is  suggested  by  Professor  Cope  that  while  in  the 
Old  World  the  intermediate  stage  between  Anchitherium  and  the 
modern  horses  was  occupied  by  this  genus,  in  the  New  World  this 
gap  was  filled  by  Protohippus.  The  true  horses  (Equus\  charac- 
terised by  the  one-toed  feet  and  the  union  of  the  anterior  inner 
column  of  the  upper  cheek-teeth  with  the  adjacent  middle  column,, 
although  now  confined  to  the  Old  World,  where  they  date  from 
the  Pliocene,  were  formerly  abundant  in  North  America  during  the 
Plistocene,  and,  as  we  have  seen,  were  likewise  represented  during 
the  same  epoch  in  South  America.  The  oldest  forms  appear  to 
be  those  from  the  Siwalik  Hills  of  northern  India ;  and  it  is  thus 
evident  that  the  group  was  originally  an  Arctogseic  one2.  The 
genus  is  now  represented  in  all  the  regions  of  eastern  Arctogaea, 
with  the  exception  of  Madagascar,  and  its  extinction  in  the  New 
World  cannot  be  satisfactorily  explained. 

1  This  pillar  forms  the  lowest  part  of  the  unshaded  area  in  figure  38. 

2  Professor  Scott,  to  whose  views  we  have  already  alluded,  in  a  paper  pub- 
lished in    Tr.  Amer.  Phil.  Soc.  Vol.  xvn.  pp.  in — 112  (1894),  is  of  opinion 
that  the  genus  Equus  was  evolved  in  North  America,  and  that  Anchitherium, 
in  its  restricted  sense,  was  off  the  direct  line.      He  arranges  the  direct  ancestral 
forms  of  the  upper  Tertiary,  from  above  downwards,  in  the  order  Protohipptis •, 
Desmatippus,  Miohippus,  and  Mesohippus ;  Anchitherium  branching  off  from 
Miohippus,  and  Hipparion  from  Protohippus. 


IV.]  PERISSODACTYLES.  169 

The  rhinoceroses  (Rhinocerotidce)  originally  had  a  distribution 
very  similar  to  that  of  the  horses,  with  the  exception  that  they 
never  entered  Neogaea;  and  they  also  agree  with  the  latter  in  their 
present  extinction  in  North  America,  where  they  are  unknown 
after  the  Pliocene.  On  both  sides  of  the  Atlantic  the  true 
rhinoceroses  appear  to  have  commenced  in  the  lower  Oligocene ; 
and  in  both  areas  the  earliest  forms  were  hornless.  Early  species 
with  a  pair  of  horns  placed  transversely  on  the  nose  are  likewise 
met  with  both  in  Europe  and  North  America;  but  the  modem 
two-horned  rhinoceroses  appear  to  be  restricted  to  the  Old  World, 
where  one  extinct  species  (Rhinoceros  antiquitatis]  ranged  as  far 
north  as  the  Arctic  circle  during  the  Plistocene  period.  On  the 
whole,  it  seems  preferable  to  include  all  the  living  and  most  of  the 
extinct  species  in  the  typical  genus  Rhinoceros  ;  the  existing  forms 


FlG.    40.       SECOND    RIGHT    UPPER    MOLAR    OF    Rhinoceros. 

A.  median  valley ;  D.  anterior,  and  E.  posterior  crests ;  F.  posterior  valley  ; 

H.  crochet. 

being  confined  to  the  Oriental  and  Ethiopian  regions.  Rhino- 
ceroses, it  may  be  observed,  differ  from  all  the  preceding  families 
of  the  suborder  by  the  upper  cheek-teeth  having  a  continuous 
outer  wall,  instead  of  being  divided  by  a  vertical  ridge  into  two 


I/O  THE   ARCTOG^IC   REALM.  [CHAP. 


distinct  lobes ;  and  while  all  the  living  forms  have  but  three  toes 
to  each  foot,  in  some  of  the  extinct  hornless  species  the  front  limb 
was  four-toed.  In  the  typical  genus  the  number  of  the  front  teeth 
is  more  or  less  reduced,  but  in  the  extinct  Hyracodon  and 
Amynodon  of  the  upper  Eocene  of  North  America  the  full  forty- 
four  teeth  were  developed ;  and  as  allied  forms  also  occur  in  the 
Oligocene,  it  would  seem  highly  probable  that  the  group  originated 
in  North  America,  whence  it  migrated  westwards  by  way  of  what 
is  now  Bering  Strait  to  attain  its  most  specialised  development  in 
the  Old  World.  It  is,  however,  noteworthy  that  the  genus 
Cadurcotherium,  from  the  French  Oligocene,  which  should  ap- 
parently find  a  place  in  this  family,  and  is  distinguished  by  the 
narrowness  of  its  upper  molars,  makes  a  curious  approximation 
in  the  structure  of  these  teeth  to  the  Neogaeic  Homalodontothe- 
rium1.  The  most  specialised  representative  of  the  family  is  the 
huge  Elasmotherium,  of  the  Siberian  Plistocene,  whose  molars  shew 
a  resemblance  to  those  of  the  Equidce. 

Another  family  of  perissodactyles  (Titanotheriidce)  is  typically 
represented  by  certain  huge,  somewhat  rhinoceros-like,  mammals, 
generally  having  a  pair  of  transversely-placed  tuberosities  on  the 
nasal  region  of  the  skull,  and  characterised  by  a  peculiar  arrange- 
ment of  the  tubercles  on  their  upper  molars.  These  teeth,  which 
have  very  short  crowns,  also  differ  from  those  of  the  rhinoceroses 
in  having  the  outer  wall  divided  into  two  lobes  by  a  vertical  ridge; 
while  the  last  lower  molar  is  distinguished  from  the  corresponding 
tooth  of  the  latter  by  having  a  third  lobe.  The  typical  genus 
Titanotherium  is  mainly  North  American,  where  it  ranges  from  the 
lower  Oligocene  to  the  upper  Eocene,  but  is  also  represented  in  the 
Tertiaries  of  the  Balkans,  although  unknown  in  those  of  western 
Europe.  An  allied  genus  (Brachydiastematotherium)  has  also  been 
recorded  from  eastern  Europe,  but  all  the  other  members  of  the 
family,  such  as  Palceosyops,  are  North  American.  This  family 
accordingly  appears  to  have  been  mainly  an  American  one,  but 
was  probably  represented  in  Asia  as  well  as  in  eastern  Europe. 

The  remarkable  genus  Chalicotherium,  whose  geological  range 
in  the  Old  World  extends  from  the  Oligocene  of  France  to  the 

1  Supra,  p.  82. 


IV.] 


PERISSODACTYLES. 


171 


THE   ARCTOG^IC    REALM.  [CHAP. 


lower  Pliocene  of  India,  while  species  also  occur  in  the  Miocene  of 
the  United  States,  has  molars  strikingly  like  those  of  the  Titano- 
theriida,  but  its  feet  differ  from  those  of  all  existing  ungulates  in 
terminating  in  huge  curved  claws  much  resembling  those  of 
the  South  American  edentates.  Indeed,  one  genus  of  the  family 
(Macrotheriuni]  was  long  regarded  as  belonging  to  the  latter 
group.  Of  the  two  genera  which  occur  in  the  European  Miocene, 
Macrotherium  has  the  fore  limb  much  longer  than  the  hinder ; 
whereas  in  Chalicotherium  (Ancylotherium)  the  two  are  more  nearly 
equal  in  length.  It  is  to  the  former  genus  that  the  North  American 
forms  are  assigned. 

The  proboscidean  ungulates,  which  differ  markedly  from  the 
two  preceding  groups  in  the  structure  both  of  their  teeth  and 
limbs,  and  are  now  represented  only  by  the  Indian  and  African 
elephant,  form  a  comparatively  small  assemblage,  most  of  whose 
members  may  be  included  in  the  family  Elephantida.  Among 
these,  the  most  specialised  types  constitute  the  genus  Elephas, 
characterised  by  the  complexity  of  the  structure  of  the  cheek-teeth, 
which  generally  assume  the  form  of  more  or  less  elevated  parallel 
plates,  with  the  intervals  filled  with  cement.  In  certain  of  the 
earlier  species  from  the  Pliocene  of  Asia  the  plates  of  these  teeth 
are,  however,  comparatively  low  and  less  numerous,  with  the 
intervening  valleys  almost  devoid  of  cement;  so  that  these 
stegodont  elephants,  as  they  are  called,  form  a  complete  transition 
towards  the  mastodons.  Commencing  in  the  Indian  Pliocene, 
elephants  ranged  over  the  whole  of  Europe  and  Asia  during  the 
Plistocene,  while  we  have  also  evidence  of  their  occurrence  during 
the  same  epoch  in  northern  Africa;  and  they  were  likewise  repre- 
sented by  two  species  in  North  America,  one  of  which  ranged  as  far 
south  as  Texas.  One  of  these  American  species  was  identical 
with  the  European  mammoth  (E.  primigenius],  while  the  second 
(E.  columbianus]  was  nearly  allied;  both  being  near  relatives  of  the 
existing  Indian  elephant.  The  extinct  stegodont  elephants  being 
confined  to  south-eastern  Asia,  it  is  interesting  to  note  that  the 
species  of  Mastodon  making  the  nearest  approximation  to  Elephas 
are  met  with  in  this  region  alone;  and  from  this  it  may  be  inferred 
that  the  evolution  of  the  latter  genus  took  place  in  that  part  of*  the 
world.  All  mastodons,  it  may  be  mentioned,  have  comparatively 


IV.] 


PROBOSCIDEANS. 


173 


simple  molar  teeth,  in  which  the  crowns  are  surmounted  by  low 
transverse  ridges,  frequently  separated  into  more  or  less  distinct 
tubercles,  and  with  the  intervening  valleys  open,  such  ridges 
varying  from  three  to  five  in  number  in  the  majority  of  the  teeth, 


FlG.    42.       LAST    UPPER    MOLAR   TOOTH    OF   A    MASTODON.       (^    nat.    size.) 

although  more  numerous  in  the  last  of  the  series.  Both  in  Europe 
and  North  America  mastodons  make  their  appearance  in  the 
Miocene,  although  in  the  latter  area  they  are  unknown  before  the 
Deep  River  beds  forming  the  upper  portion  of  that  stage;  and 
whereas  they  disappeared  in  the  Old  World  with  the  Pliocene, 
they  persisted  in  the  New  till  the  succeeding  Plistocene  age. 
During  the  Pliocene,  as  we  have  seen  in  the  last  chapter,  they 
obtained  an  entrance  into  South  America,  so  that  they  cannot  be 
regarded  as  absolutely  characteristic  of  Arctogaea.  On  the  whole, 
it  is  probable  that  the  group  originated  in  the  Old  World,  although 
we  are  still  in  the  dark  as  to  its  relationship  to  other  ungulates. 

The  only  other  Arctogaeic  genus  of  proboscideans  is  Dino- 
therium,  which  constitutes  a  family  by  itself,  and  is  known  from  the 
Miocene  and  Pliocene  of  Europe  and  India,  but  is  unrepresented 
in  America.  In  these  animals  only  one  of  the  true  molars  has 
three  ridges,  the  others  having  but  two;  and  tusks  were  present  in 
the  lower  jaw  alone. 

A  fourth  subordinal  group  of  the  ungulates,  which  is  more 
primitive  than  any  of  the  foregoing,  and  has  been  designated  by 
Professor  Cope  the  Amblypoda,  or  short-footed  group,  has  one 


THE   ARCTOG^IC    REALM.  [CHAP. 

family  (Coryphodontidce]  common  to  the  lower  Eocene  of  both 
hemispheres,  while  a  second  family  ( Uintatheriid<z),  of  later  age  is 
strictly  North  American.  All  these  animals  had  limbs  of  an 
elephantine  type,  each  foot  being  furnished  with  five  toes,  and  the 
bones  of  the  wrist  and  ankle  joints  arranged  on  the  linear  plan. 
The  genus  Coryphodon,  which  includes  species  varying  in  size 
from  a  tapir  to  a  rhinoceros,  had  forty-four  teeth,  with  the  canines 


FIG.  43.     LEFT  UPPER  CHEEK-TEETH  OF   Coryphodon ;  reduced. 

well  developed,  and  the  molars  bearing  prominent  oblique  ridges. 
First  discovered  in  the  London  Clay,  and  subsequently  in  the 
lower  Eocene  of  France,  this  genus  has  since  been  recognised  in 
the  lower  Eocene  of  the  United  States,  where  nearly  perfect 
skeletons  have  been  obtained. 

In  a  still  more  primitive  group  known  as  the  Condylarthra, 
which  apparently  contains  the  ancestral  stock  of  both  the  artio- 
dactyles  and  perissodactyles,  it  is  believed  that  the  genus  Phena- 
codus  (the  ultimate  parent  of  the  horses),  typically  occurring  in  the 
Wahsatch  Eocene  of  the  United  States,  is  represented  in  the  Swiss 
Eocene;  and  the  same  has  been  stated  to  be  the  case  with 
Protogonia. 

Summarising  the  results  of  the  foregoing  survey,  it  may  be 
stated  that  as  regards  its  mammalian  fauna  Arctogaea 

Summary  of  . 

the  character-  as  a  whole  is  characterised  by  the  following  features, 
mammalian  Absence  of  monotremes  and  diprotodont  marsu- 
faunaof  Arcto-  pials.  No  existing  polyprotodont  marsupials  except 
opossums,  and  these  only  in  its  western  half.  No 
Tertiary  marsupials  known  except  opossums,  although  other  types 
probably  existed  in  South-eastern  Asia.  No  existing  edentates1, 
and  fossil  ones  only  in  North  American  Plistocene  and  Pliocene. 
No  marmosets  (Hapalida),  and  no  monkeys  of  the  family  Cebidce. 
Bats  of  the  family  Phyllostomatida  wanting,  save  for  a  few  on  the 
1  The  aard-varks  and  pangolins  are  here  excluded  from  the  Edentata. 


IV.]  LIST   OF   FAMILIES.  1/5 

Pacific  side  of  North  America.  Insectivores  abundant.  In  the 
following  list  of  more  or  less  widely  distributed  families  the  ma- 
jority are  for  the  most  part  confined  to  this  realm,  while  the  others 
contain  exclusively  Arctogaeic  genera.  Such  groups  as  are  prac- 
tically confined  to  Arctogaea  are  printed  in  italics,  those  which  are 
extinct  having  an  *  prefixed ;  while  the  letter  H.  denotes  such  of 
the  existing  ones  as  are  restricted  to  the  Holarctic  region  : — 

INSECTIVORA. 

Talpida.     H. 

Soricidce.     Enters  Austro-Malayan  region  and  Mexican 

sub-region. 
CARNIVORA. 

*  Hycenodoriidce. 

*  Proviverridce. 

*  ArctocyonidcE. 

RODENTIA. 

Sciuridae.     Mainly  Arctogseic,  although  species  of  Sciurus 

extend  in  Neogaea  as  far  south  as  Paraguay. 
Dipodidce. 
Castorida.     H. 

*  Theridomyida.     H. 
Lagomyidce.     H. 

UNGULATA. 

*  Chceropotamidtz. 

*  A  nthracotheriida. 

*  Ccenotheriidce. 

TraguhdcE.     Extinct  in  N.  America. 

Cervidae.     The  genus  Cervus  exclusively  Arctogaeic. 

Bovida.    Represented  by  Bos  depressicornis  in  Celebes. 

*  Lophiodontidce. 

*  Palaotheriidce. 

Rhinocerotida.     Extinct  in  W.  Arctogaea. 

*  Chalicotheriidce. 

*  Titanotheriida*.     Mainly    W.    Arctogaeic    but    occurring 

in  the   Balkans. 
Elephantidae.     Elephas. 

*  CoryphodontidcK. 


THE    ARCTOG/EIC    REALM. 


[CHAP. 


f  ATUL TITUBERCULA  TA. 

*  Plagiaulacidce. 

* Bolodontidce.     Only  Secondary  in  Eastern  Arctogsea. 

The  following  table  exhibits  some  of  the  better  known  Tertiary 
mammalian  genera  common  to  both  halves  of  Arctogaea,  together 
with  allied  types  restricted  to  the  western  and  eastern  hemispheres ; 
such  as  are  still  existing  being  indicated  by  a  f : — 

W.   HEMISPHERE.          BOTH  HEMISPHERES.  E.  HEMISPHERE. 


LEMUROIDEA. 

Anaptomorphus. 
CARNIVORA. 


Hyopsodus. 


Adapis. 
Microchoerus. 


(Also 


Machasrodus 
Neogaea  in  Plistocene). 


Nimravus. 

Archaelurus. 

Temnocyon. 

Hyaenocyon. 

Daphaenus. 


Eusmilus. 


Claenodon. 
RODENTIA. 


UNGULATA. 

Achaenodon. 

Homacodon. 
Procamelus,  etc. 


Hyaenodon. 
Pterodon. 
Oxysena. 
Proviverra. 


Chalicomys. 
Plesiarctomys. 

Elotherium. 
Ancodus. 


^Elurictis. 


Amphicyon. 

Dinocyon. 

Hyaenarctus. 


Arctocyon. 


Chceropotamus. 

Anthracotherium. 

Dichobunus. 

Caenotherium. 
t  Camelus. 
f  Dorcatherium. 


IV.] 


WESTERN    AND   EASTERN    FAUNAS. 


177 


UNGULATA  (continued). 
Leptomeryx. 
Hypertragulus. 


Colodon. 
Systemodon. 


Protohippus. 


Hyracodon. 
Amynodon. 


Palaeosyops. 


Prodremotherium. 

Protapirus. 

Helaletes.  Lophiodon. 

Hyracotherium. 

Pachynolophus. 

Hyrachyus. 

Palaeotherium. 
Anchitherium.     Anchilophus. 

Hipparion. 

t  Equus  (also  Neogaeic). 
t  Rhinoceros. 

Cadurcotherium. 

Elasmotherium. 
Titan  otherium. 

Brachydiastematotherium. 
Chalicotherium.  Macrotherium. 
t  Elephas. 
Mastodon  (also  Neogaeic). 

Dinotherium. 
Coryphodon. 

( Typically  N.  American, 
Phenacodus  IV*        J      , 

i      but  stated  also  to 
Protogoma    i  .     ^ 

I     occur  in  Europe. 


In  the  foregoing  table,  only  some  of  the  better  known  types 
have  been  selected,  but  these  suffice  to  show  that  throughout  the 
Tertiary  epoch  a  certain  number  of  genera  were  common  to 
western  and  eastern  Arctogaea.  It  is  true  that,  with  the  exception 
of  those  still  existing,  we  have  no  evidence  that  any  of  these  ever 
reached  the  Ethiopian  region ;  and  it  is  quite  probable  that  many 
or  the  whole  of  them  never  did  so.  By  its  present  fauna  that 
region  is,  however,  so  closely  connected  with  the  Pliocene  and 
modern  mammals  of  Asia  and  Europe,  that  there  can  be  no  ques- 
tion of  its  right  to  inclusion  in  the  same  realm ;  and  this  being  so, 
L.  12 


1/8  THE   ARCTOG^EIC   REALM.  [CHAP.  IV. 

Madagascar  cannot  be  excluded.  Still,  it  is  quite  probable  that 
during  the  later  Tertiary  epoch  the  Ethiopian  and  Malagasy 
regions  were  almost  as  distinct  from  the  rest  of  Arctogaea  as  are 
Neogaea  and  Notogaea  at  the  present  day,  and  if  such  conditions 
had  continued,  the  former  areas  would  have  been  entitled  to  con- 
stitute a  realm  by  themselves. 

In  the  sequel  we  shall  discuss  the  whole  number  of  existing 
genera  of  non-volant  terrestrial  mammals  common  to  the  eastern 
and  western  halves  of  the  Holarctic  region,  and  likewise  such 
living  and  extinct  types  as  are  respectively  peculiar  to  Eastern  and 
Western  Arctogsea.  These,  taken  in  conjunction  with  the  fore- 
going tables,  will  show  that,  in  spite  of  the  forms  common  to  the 
two  latter  areas,  there  has  always  been  a  large  number  of  types 
restricted  to  one  side  or  the  other  of  the  Atlantic  basin.  And 
this  leads  to  the  conclusion  that,  although  during  a  considerable 
portion,  or  the  whole  of  the  Tertiary  period,  there  was  a  free  land- 
communication  between  North  America  and  Eastern  Asia  by  way 
of  Bering  Strait,  yet  that  this  connecting  land  must  have  been 
comparatively  narrow,  so  that  the  faunas  of  the  more  southern 
portions  of  both  areas  developed  to  a  great  extent  independently 
of  one  another. 

Not  the  least  curious  feature  in  connection  with  the  com- 
munity of  types  on  the  two  sides  of  the  Atlantic  is  the  precise 
parallelism  in  the  development  of  many  of  the  groups  in  both 
areas.  In  both,  for  instance,  the  phyla  of  the  horses  and  rhino- 
ceroses were  practically  similar,  although  it  is  thought  that  the 
stage  occupied  in  the  one  area  by  Hipparion  was  held  in  the  other 
by  Protohippus.  If  this  particular  suggestion  should  prove  to  be 
well  founded,  it  will  be  self-evident  that  the  true  horses  have  been 
independently  evolved  in  the  two  areas ;  and  it  almost  seems  as 
if  the  same  had  been  the  case  with  the  rhinoceroses  and  certain 
other  groups.  Had  the  culminating  forms  been  devolved  in  only 
one  of  the  two  areas,  we  should  not  expect  to  find  the  whole  of 
the  ancestral  links  present  in  both. 


CHAPTER   V. 


EASTERN   ARCTOG^EA. 

Mammalian  groups  peculiar  to  Eastern  Arctogsea  —  Tertiary  Mammalian 
Faunas  of  Eastern  Arctogaea  —  Oligocene  Fauna  —  Miocene  Fauna  —  Older 
Pliocene  Fauna  —  Pikermi  and  allied  Faunas  —  Siwalik  Fauna  —  Higher 
Pliocene  Faunas. 

ALTHOUGH  northern  Europe  and  Asia  forms  but  one  zoological 
region  with  the  corresponding  part  of  North  America, 
yet  there  are  numerous  groups  of  mammals  con- 


fined  respectively  to  its  eastern  and  western  portions,  ^u>  Eastern 
which  clearly  show  that  the  communication  between 
the  two  areas  was  always  more  or  less  limited.  In  this  chapter 
attention  will  be  first  directed  to  some  of  the  most  striking 
peculiarities  of  the  mammalian  fauna  of  Eastern  Arctogaea,  after 
which  the  whole  fossil  fauna  may  be  taken  into  consideration. 

In  addition  to  the  total  absence  of  existing  opossums  (Didel- 
phyid(E\  and  the  presence  in  its  warmer  portions  of  fruit-bats 
(Pteropodidce),  which,  however,  are  common  to  Notogaea,  Eastern 
Arctogaea  is  especially  characterised  as  being  the  home  of  all  the 
higher  Primates;  namely  the  family  Stmiidce,  which  includes  the 
man-like  apes  and  gibbons,  and  the  Cercopithecidcz,  embracing  all 
the  other  Old  World  monkeys.  From  the  South  American  monkeys 
(Cebidcz)  both  these  families  are  broadly  distinguished  by  having 
two  pairs  of  premolar  and  three  of  molar  teeth,  whereas  in  the 
former  group  there  are  three  pairs  of  both  premolar  and  molar 
teeth.  Not  only  are  the  two  families  in  question  confined  at  the 
present  day  to  the  Eastern  hemisphere,  but  the  same  appears  to 
have  been  the  case  at  all  epochs,  since  no  trace  of  a  fossil  monkey 
has  ever  been  recorded  from  North  America.  This  remarkable 
isolation  of  the  distributional  areas  of  the  Simiidcz  and  Cercopi- 
thecidce  on  the  one  hand,  and  of  the  Cebidce.  (and  Hapalidcz)  on 

12  —  2 


ISO  EASTERN   ARCTOG^A.  [CHAP. 

the  other,  points  unmistakeably,  in  spite  of  their  external  similarity, 
to  the  dual  origin  of  the  monkeys  of  the  Old  and  New  Worlds. 
Both  may,  however,  have  originated  from  different  groups  of 
lemuroids,  which,  as  indicated  in  an  earlier  chapter,  ranged  over 
the  whole  of  Northern  Arctogaea  during  the  earlier  part  of  the 
Tertiary  epoch. 

At  the  present  day  the  man -like  apes,  which  are  few  in 
number,  have  an  extremely  limited  distribution.  The  chim- 
panzees (Anthropopithecus)  are  restricted  to  Equatorial  Africa,  the 
gorilla  (Gorilla)  is  found  only  in  the  hottest  regions  of  Western 
Africa,  the  orangs  (Simla}  are  confined  to  the  islands  of  Borneo 
and  Sumatra,  and  the  smaller  gibbons  (Hylobates)  are  inhabitants 
of  South-eastern  Asia,  from  Assam  and  Burma  to  Hainan.  Ex- 
tinct species  of  chimpanzees  and  orangs  occur,  however,  in  the 
Pliocene  of  Northern  India;  while  gibbons  are  met  with  in  the 
Miocene  of  France  and  Baden,  although  there  is  some  difference 
of  opinion  whether  these  are  generically  identical  with  the  Asiatic 
forms,  or  should  be  assigned  to  a  genus  apart  (Pliopithecus}.  The 
former  deposits  have  also  yielded  remains  of  a  large  extinct  ape 
(Dryopithecus))  apparently  of  a  somewhat  more  generalised  type 
than  all  the  existing  forms. 

The  ordinary  monkeys  and  apes  (Cercopithecidce)  have  a  wider 
distribution,  ranging  over  most  of  the  warmer  parts  of  Eastern 
Arctogaea,  and  being  represented  by  a  single  species  at  Gibraltar, 
and  by  two  others  in  Moupin,  in  Eastern  Tibet,  and  a  fourth  in 
Japan.  This  family,  by  the  way,  is  not  exclusively  Arctogaeic, 
since,  as  we  have  seen,  one  species  of  a  peculiar  genus  (Cynopi- 
thecus)  inhabits  Celebes.  Apart  from  the  latter,  the  family  is 
represented  by  eight  living  genera,  among  which  the  Ethiopian 
Papio  has  extinct  representatives  in  the  Pliocene  and  Plistocene 
of  India,  as  have  also  the  Asiatic  Macacus  and  Semnopithecus.  The 
two  latter  genera  also  occur  in  the  Pliocene  of  France  and  Italy; 
and  a  tooth  of  a  species  of  Macacus  has  been  obtained  from  the 
Plistocene  brick-earths  of  Essex.  In  addition  to  these,  there  are 
certain  extinct  genera  from  the  European  Tertiaries;  among 
which  Mesopithecus  from  the  lower  Pliocene  of  Greece  agrees  with 
Macacus  in  its  short  and  stout  limbs,  but  approximates  to 
Semnopithecus  in  the  character  of  its  skull  and  dentition.  Dolicho- 


V.]  DISTINCTIVE   MAMMALS.  l8l 

pithecus,  from  the  French  Pliocene,  has  a  longer  muzzle;  while 
Oreopithecus,  from  the  Italian  Miocene,  seems  to  connect  the 
CercopitheddcB  with  the  Simiida. 

It  is  thus  evident  that  during  the  latter  portion  of  the  Tertiary 
epoch  monkeys  and  apes  were  spread  over  the  greater  part  of 
Eastern  Arctogaea;  and  their  extensive  diffusion  is  a  proof  that 
this  half  of  the  realm  could  only  have  been  connected  with  North 
America  by  land  situated  so  far  north  that  it  formed  an  impass- 
able barrier  to  these  animals.  Although  the  smaller  extinct 
European  monkeys  do  not  necessarily  indicate  a  very  high 
temperature  in  the  regions  they  inhabited,  there  can  be  little  doubt 
that  at  the  era  when  Dryopithecus  flourished  southern  Europe  at 
least  enjoyed  a  moist  tropical  climate. 

Not  less  characteristic  of  Eastern  Arctogaea  are  the  existing 
lemuroids,  of  which  there  are  three  families ;  the  largest  (Lemurida) 
ranging  over  the  Oriental,  Ethiopian,  and  Malagasy  regions,  the 
Tarsiidce,  with  a  single  genus,  being  exclusively  Oriental,  while 
the  sole  representative  of  the  Ckiromyida  is  Malagasy.  The 
numerous  existing  members  of  the  Lemuridce.  are  all  characterised 
by  the  first  of  the  three  lower  premolar  teeth  assuming  the  form 
and  functions  of  a  canine;  and  as  this  feature  is  unknown  in 
any  of  the  Tertiary  representatives  of  the  sub-order,  this  family 
appears  to  be  exclusively  confined  to  the  area  under  consideration. 
For  the  most  part,  the  Oligocene  lemuroids  of  Europe  seem 
likewise  to  have  been  markedly  distinct  from  those  of  North 
America.  Microchcerus^ ,  for  instance,  which  is  represented  both 
in  France  and  England,  indicates  a  family  characterised,  among 
other  features,  by  the  general  presence  of  only  three  pairs  of 
premolar  teeth  in  each  jaw ;  and  Adapts,  which  is  likewise 
common  to  the  same  two  countries,  has  four  such  teeth. 

Although  there  are  several  families  of  Insectivora  peculiar  to 
the  eastern  hemisphere,  the  only  one  of  these  with  a  wide  distri- 
bution is  that  of  the  hedgehogs  and  their  allies  (Erinaceida),  which 
has  representatives  in  the  eastern  Holarctic,  Oriental,  and  Ethiopian 
regions.  Although  none  are  known  from  America,  extinct  repre- 
sentatives of  this  family  are  common  in  the  European  Oligocene. 

1  Teeth  figured  on  p.  156. 


1 82  EASTERN    ARCTOG^EA.  [CHAP. 

Of  these  Pal&oerinaceus  appears  to  be  allied  to  the  true  hedgehogs 
(Erinaceus) ;  whereas  other  genera,  such  as  Necrogymnurus, 
connect  the  former  with  the  existing  long-tailed  and  spineless 
Gymnura  of  the  Malayan  islands.  This  group  is  thus  essentially 
characteristic  of  Eastern  Arctogaea  as  a  whole. 

Turning  to  the  Carnivora,  we  have,  in  addition  to  the  Prote- 
leidce,  of  which  the  sole  representative  is  the  African  aard-wolf,  two 
important  families  practically  confined  to  this  half  of  the  realm. 
The  first  of  these  is  the  extensive  group  of  the  civets,  mungooses, 
and  their  allies  ( Vtverridce),  which  has  no  representatives  in  the 
New  World,  although  a  single  species  in  two  genera  ranges  into 
the  Austro-Malayan  region.  Out  of  a  total  of  twenty-three  genera 
included  in  this  family  only  one  mungoose  (Herpestes)  and  the 
common  genet  (Genetta)  range  into  Europe,  most  of  the  other 
forms  being  confined  to  the  Oriental,  Ethiopian,  and  Malagasy 
regions.  Civets  (  Viverra)  and  ichneumons,  together  with  several 
remarkable  extinct  genera,  such  as  Stenoplesictis,  were,  however, 
common  in  the  European  Tertiaries,  from  the  lower  Oligocene 
upwards.  And  from  the  circumstance  that  the  last-named  genus 
presents  characters  connecting  the  Viverridce  with  the  weasel 
tribe  (Mustelida),  it  would  seem  probable  that  the  latter  family 
was  originally  evolved  in  Eastern  Arctogaea,  although  it  has  now  a 
considerable  number  of  American  representatives. 

By  means  of  certain  extinct  forms  from  the  lower  Pliocene  of 
Southern  Europe  and  Northern  India  known  as  Ictitherium,  the 
civets  are  very  closely  connected  indeed  with  the  hyaenas 
(ffy&nidce) ;  the  three  living  representatives  of  which  may  be 
included  in  the  single  genus  Hyana.  Although  the  striped  hyaena 
(H.  striata)  is  now  confined  to  Southern  Asia  and  Northern 
Africa,  it  occurred  in  the  Pliocene  epoch  in  France  and  England; 
while  the  larger  spotted  hyaena  (ff.  crocutd]  of  Southern  Africa 
ranged  during  the  Plistocene  era  over  the  greater  part  of  temperate 
Europe,  and  likewise  extended  eastwards  as  far  as  India. 
Numerous  extinct  species  of  the  same  genus  are  found  in  the 
Pliocene  of  Europe  and  India;  and  two  extinct  genera  from  the 
same  deposits — Hyanictis  and  Palhyizna — connect  the  living 
forms  with  the  aforesaid  Ictitherium.  Although  one  extinct 
Tertiary  North  American  genus  has  been  tentatively  assigned  to 


V.]  RODENTS.  183 

it,  the  family  is  thus  essentially  an  Eastern  Arctogaeic  one;  and  it 
may  be  assumed  that,  as  its  living  representatives  are  inhabitants  of 
hot  climates,  the  extinct  forms  were  unable  to  exist  sufficiently  far 
north  to  permit  them  to  cross  by  the  bridge  of  land  via  Bering 
Strait. 

Among  the  rodents,  in  addition  to  the  two  widely  spread 
families — Myoxidcz  and  Spaladdce. — confined  to  Eastern  Arctogaea, 
we  find  in  the  Muridce  the  sub-family  of  the  gerbils  (Gerbillina) 
similarly  restricted,  their  range  including  the  whole  of  Eastern 
Arctogaea  with  the  exception  of  the  Malagasy  region. 

The  subfamily  Murtna,  which  includes  the  true  rats  and  mice 
(Afus)  is  likewise  restricted  to  the  Old  World.  These  rodents 
differ  from  the  hamsters  (Cricetus)  and  the  New  World  white- 
footed  mice  (Sitomys),  which,  with  other  forms,  constitute  the 
sub-family  Cricetince,  by  the  molar  teeth  in  the  upper  jaw  having 
their  tubercles  arranged  in  three  longitudinal  rows ;  whereas  in  the 
latter  they  form  only  a  double  series.  Distributed  over  all  the 
regions  of  Eastern  Arctogaea,  with  the  exception  of  Madagascar, 
this  group  is  also  represented  in  the  Australian  region.  Of  the 
two  extensively  distributed  families  restricted  to  the  area  under 
consideration,  the  first  is  that  of  the  dormice  (Myoxidce)  whose  range 
includes  the  Eastern  Holarctic  and  Ethiopian  regions.  Distin- 
guished from  all  other  rodents  by  the  absence  of  a  caecum  or 
blind  appendage  to  the  intestine,  and  further  characterised  by 
the  complicated  infoldings  of  the  enamel  on  the  crowns  of  their 
molar  teeth,  these  beautiful  little  creatures  now  attain  their 
maximum  development  in  Africa.  In  a  fossil  state  they  are  first 
known  from  the  lower  Oligocene  of  Europe,  and  are  likewise 
common  in  the  Miocene.  Another  family  which  does  not  range 
beyond  the  limits  of  Eastern  Arctogaea  is  that  of  the  Spalacida, 
typically  represented  by  the  great  mole-rat  (Spalax  typhlus] — a 
blind  creature,  ranging  over  south-eastern  Europe,  Persia,  Meso- 
potamia, Syria,  and  Egypt.  The  allied  genus  (Rhizomys),  in  which 
the  eyes,  although  minute,  are  not  covered  with  skin,  includes 
several  species,  whose  distributional  area  embraces  the  north  of 
India,  Tibet,  China,  Burma,  Malaysia,  and  Abyssinia  ;  but  the 
three  remaining  genera  are  restricted  to  the  Ethiopian  region. 
The  whole  of  the  foregoing  belong  to  the  mouse-like,  or  Myo- 


1 84  EASTERN    ARCTOGvEA.  [CHAP. 

morphous  section  of  the  order,  but  among  the  Hystricomorphous 
rodents  we  have  the  typical  porcupines  (Hystricince),  in  which  the 
tail  is  never  prehensile,  practically  confined  to  Eastern  Arctogsea, 
where  they  range  over  south-eastern  Europe,  and  the  Ethiopian 
and  Oriental  regions.  The  Javan  species  of  the  typical  genus 
(Hystrix  javanicd]  is,  however,  found  in  the  island  of  Timor, 
in  the  Austro-Malayan  region,  although  it  is  doubtless  of  late 
introduction  there,  and  may  not  improbably  have  been  transported 
by  human  agency.  In  a  fossil  state  porcupines  of  this  sub-family 
are  common  in  the  European  Tertiary  as  far  down  as  the  lower 
Oligocene. 

Turning  to  the  ungulates,  we  have  in  the  artiodactyle  section 
two  closely  allied  families,  which — if  we  except  certain  pigs  from  the 
Austro-Malayan  region  and  Papua,  which  may  have  been  originally 
introduced  by  man — are  restricted  to  the  area  under  considera- 
tion. Both  these  families,  moreover,  have  representatives,  either 
living  or  extinct,  in  all  the  regions  of  Eastern  Arctogsea,  inclusive 
even  of  Madagascar,  so  that  they  may  be  reckoned  among  its  most 
characteristic  mammals.  The  Hippopotamidtz — now  restricted  to 
the  Ethiopian  region,  where  they  are  represented  by  the  widely- 
spread  common  Hippopotamus  amphibius,  and  the  much  smaller 
terrestrial  H.  liberiensis  of  the  West  Coast — ranged  during  the 
Plistocene  and  upper  Pliocene  epochs  over  the  greater  part  of 
Europe  as  far  north  as  England ;  one  species  from  these  deposits 
being  apparently  indistinguishable  from  the  common  African  form. 
Extinct  species  are  met  with  in  the  Plistocene  of  Algeria,  the 
Plistocene  and  Pliocene  of  India,  the  Pliocene  of  Burma,  and  the 
superficial  deposits  of  Madagascar;  some  of  these  differing  from 
the  common  hippopotamus  by  the  presence  of  three,  in  place  of 
two,  pairs  of  incisor  teeth  in  each  jaw.  Whatever  may  have  been 
the  case  with  the  swine,  it  is  evident  that  the  hippopotami  were 
never  able  to  exist  sufficiently  far  north  to  cross  by  way  of  Bering 
Strait  into  the  New  World.  In  the  pigs  (Suidez) — which  among 
other  features  differ  from  the  Hippopotamidce  by  the  nostrils  being 
perforated  in  a  fleshy  disc  at  the  extremity  of  the  muzzle,  and  like- 
wise by  the  structure  of  the  teeth — the  typical  group  of  the  genus 
Sus  ranges  over  most  of  the  Eastern  Holarctic  and  the  whole 
of  the  Oriental  region,  being  replaced  in  the  Ethiopian  and 


V.]  UNGULATES.  185 

Malagasy  regions  by  the  potamochcerine  group,  frequently  reckoned 
as  a  distinct  genus.  The  Ethiopian  region  is,  however,  the  sole 
habitat  of  the  wart-hogs  (Phacochoerus}. 

Among  extinct  artiodactyles  we  have  two  well-marked  families, 
distinguished  from  the  foregoing  by  the  crescentic  columns  of 
their  short-crowned  cheek  teeth — the  Anoplotheriidce  and  Dicho- 
dontidce — likewise  confined  to  Eastern  Arctogaea,  although  their 
remains  have  hitherto  been  obtained  only  from  the  eastern  section 
of  the  Holarctic  region.  The  first  of  these  includes  several  genera 
from  Oligocene  strata,  characterised  by  having  three  columns  on 
the  front  half,  and  two  on  the  hinder  half  of  their  upper  molars. 
In  the  typical  Anoplotherium  there  were  forty-four  teeth,  arranged 


FlG.    44.        LAST    FIVE   RIGHT    UPPER  CHEEK-TEETH    OF   AN    Anoplotherium. 

in  a  continuous  even  series,  and  the  feet  were  provided  with  either 
three  or  two  toes.  Dacrytherium  differs  by  the  molars  being  more 
like  those  of  Ancodus,  and  also  by  the  deep  cavity  for  the 
reception  of  a  gland  on  each  side  of  the  face  in  front  of  the  eye ; 
while  the  small  and  elegantly  formed  animals  described  as  Xiph- 
odon  have  the  crowns  of  the  first  three  premolar  teeth  elongated 
and  trenchant,  the  feet  being  two-toed.  In  the  Dichodontida,  of 
which  there  are  likewise  several  genera,  the  cheek-teeth  are  more 
completely  selenodont,  with  only  four  columns  on  the  crowns  of 
the  upper  molars ;  and  it  is  not  improbable  that  in  this  family  we 
have  the  ancestral  types  of  both  the  chevrotains  and  the  deer. 

In  the  Camelidce,  as  we  have  already  seen,  the  typical  genus 
Camelus,  which  is  found  living  (although  not  in  a  wild  state)  in  the 
Eastern  Holarctic,  Oriental,  and  Ethiopian  regions,  and  fossil  in 
the  Plistocene  of  Algeria  and  the  Pliocene  of  India,  is  likewise 
confined  to  Eastern  Arctogsea.  And  the  same  is  true,  both  in  the 


1 86  EASTERN    ARCTOG^A.  [CHAP. 

recent  and  fossil  state,  with  the  genera  Tragulus  and  Dorcatherium, 
which  at  the  present  day  alone  represent  the  Tragnlida. 

The  family  of  giraffes  (Giraffidce),  of  which  the  Ethiopian 
Giraffa  cameiopardalis  is  the  sole  existing  survivor,  was  formerly 
extensively  distributed  over  the  area  under  consideration,  to  which 
it  appears  to  have  been  always  restricted,  albeit  represented  by 
a  considerable  number  of  generic  types.  True  giraffes  (Giraffd) 
ranged  during  the  Pliocene  epoch  over  Greece,  Persia,  India, 
and  China,  and  allied  types  are  to  be  found  in  Visnutherium 
of  the  Pliocene  of  India  and  Burma,  and  Helladotherium  from 
the  corresponding  formation  of  Greece.  Still  more  gigantic  than 
the  latter  were  the  huge  Hydaspitherium,  Bramatherium,  and 
Sivatherium,  of  the  Indian  Pliocene,  in  all  of  which  the  simple 
horns  of  the  giraffes  were  replaced  by  large  antler-like  appendages, 
differing  considerably  in  their  arrangement  in  the  different  genera. 
Other  members  of  the  family  are  Samotherium,  of  the  Pliocene  of 
the  Isle  of  Samos,  and  Palceotragus  from  the  equivalent  deposits 
of  Attica,  in  both  of  which  the  females  appear  to  have  been  horn- 
less, although  the  males  had  a  pair  of  simple,  compressed,  and 
nearly  upright  horn-cores.  The  former  is  represented  by  a  species 
rivalling  the  giraffe  in  the  size  of  its  skull,  but  the  latter  was  a 
much  smaller  animal.  This  group  likewise  extended  to  Northern 
Africa,  where  a  large  species  from  the  Algerian  Pliocene  has  been 
described  under  the  name  of  Libytherium. 

Although  the  extensive  family  of  the  Bovidce,  including  the 
oxen,  sheep,  goats,  antelopes,  etc.,  is  now  represented  in  the 
northern  part  of  the  western  Holarctic  region  by  the  American 
bison  (Bos  americanus\  the  bighorn  (Ovis  canadensis],  the  musk- 
ox  (Ovibos  moschatus],  and  the  so-called  Rocky  Mountain  goat 
(Haploceros  montanus),  together  with  a  few  extinct  forms  from  the 
superficial  deposits,  while  the  anoa  (Bos  depressicornis)  is  peculiar 
to  Celebes,  the  greater  number  of  its  representatives  belong  to 
Eastern  Arctogsea.  The  whole  of  the  numerous  genera  of  ante- 
lopes, together  with  the  true  goats,  as  well  as  the  great  majority  of 
the  sheep,  are,  for  instance,  restricted  to  this  area.  Moreover, 
whereas  the  musk-ox  is  now  solely  North  American,  it  was  common 
in  Europe  during  the  Plistocene ;  while  the  bighorn  is  closely 
allied  to  the  Kamschatkan  wild  sheep  (O.  nivicola),  and  the 


V.]  EFFODIENTIA.  l8/ 

American  bison  not  far  removed  from  its  Caucasian  cousin  (Bos 
bison],  so  that  all  these  forms  are  probably  descended  from 
ancestors  inhabiting  Eastern  Arctogaea. 

In  the  perissodactyle  section  of  the  ungulates,  if  we  take  fossil 
forms  into  account,  there  are  no  families  peculiar  to  this  area;  but 
among  extinct  forms  we  have  the  large  Oligocene  genus  Palceo- 
therium1,  and  the  Eocene  Lophiodon  absolutely  restricted  to  it. 

Although  occurring  only  in  Syria  and  the  Ethiopian  region,  and 
at  present  unknown  in  a  fossil  state,  the  peculiar  subordinal  group 
of  ungulates  represented  solely  by  the  hyraces  (Procaviidce)  per- 
haps deserves  mention  among  the  types  characteristic  of  Eastern 
Arctogaea.  A  nearly  similar  observation  applies  to  the  extinct 
proboscidean  family  Dinotheriidce,  in  which  the  single  known 
genus  (Dinotherium)  ranges  from  the  Miocene  and  Pliocene  of 
Europe  to  the  Pliocene  of  Northern  India. 

Of  the  edentates,  with  the  exception  of  certain  very  doubtful 
forms  from  the  French  phosphorites,  which  may  prove  to  be 
reptilian,  we  have  no  evidence  of  the  existence  of  any  representa- 
tives in  the  Old  World.  There  are,  however,  in  Eastern  Arcto- 
g?ea  two  very  peculiar  families  commonly  assigned  to  the  same 
order  as  the  latter,  although  it  seems  preferable  to  regard  them  as 
indicating  an  ordinal  group  (Effodientia)  by  themselves.  Of  these 
the  pangolins  (Manidce),  which  are  distinguished  from  all  other 
mammals  by  their  covering  of  overlapping  horny  scales,  are  now 
confined  to  the  Oriental  and  Ethiopian  regions,  to  which  the  one 
living  genus  Manis  is  common ;  but  they  appear  to  have  been 
represented  in  the  Oligocene  phosphorites  of  France  by  smaller 
extinct  forms,  to  which  the  names  Necromanis  and  Leptomanis* 
have  been  given.  The  second  family,  Orycteropodidce,  of  which 
the  only  living  members  are  the  Ethiopian  aard-varks  (Orycteropus), 
differ  very  widely  from  the  last,  the  body  being  nearly  naked,  and 
the  molar  teeth  characterised  by  a  peculiarly  complex  structure 
which  is  unique  in  the  whole  mammalian  class.  A  fossil  species 
of  the  existing  genus  has  been  discovered  in  the  lower  Pliocene  of 
Samos  and  Maraga,  in  Persia ;  while  the  extinct  genus  Pal&orycte- 

1  Teeth  figured  on  p.  166. 

2  The  so-called  Palceomanis,  from  the  Pliocene  of  Samos,  turns  out  to  have 
been  founded  on  remains  of  an  ungulate. 


1 88  EASTERN   ARCTOG^EA.  [CHAP. 

ropus,  from  the  French  phosphorites,  is  believed  to  have  belonged 
to  the  same  family,  so  that  both  groups  appear  formerly  to  have 
been  widely  distributed. 

Summarising  the  results  of  the  foregoing  survey,  we  may  put 
in  a  tabular  form  the  leading  features  of  the  mammalian  fauna  of 
Eastern  as  distinct  from  that  of  Western  Arctogsea.  In  the  sub- 
joined table  the  letters  E.,  M.,  O.,  H.  respectively  indicate  the 
Ethiopian,  Malagasy,  Oriental,  and  Eastern  Holarctic  regions ; 
and  when  a  family  is  represented  in  any  of  such  regions  only  in  a 
fossil  state,  a  f  is  added  to  the  denoting  letter.  The  names  of  such 
families  or  groups  as  are  practically  peculiar  to  the  area 'under 
consideration  are  printed  in  italic  type  ;  while  extinct  groups  have 
an  *  prefixed. 

PRIMATES. 

Simiidce.     O.  E.  H.f 

Cercopithecidcz.     O.  E.  H. ;  also  extending  into  the  Austro- 

Malayan  region. 
Lemuridtz.     O.  E.  M. 
*  Microchoeridce.     H. 
* Adapida.     H. 

CHIROPTERA. 

Pteropodidse.     O.  E.  ;  also  common  to  Notogaea. 
INSECTIVORA. 

Erinaceidce.     O.  E.  H. 

CARNIVORA. 

Viverridce.     O.  E.  H.  M.;    two  species   extending  their 

range  into  the  Austro- Malayan  region. 
Hycenidcz.     O.  E.  H.f 

RODENTIA. 

Muridae  ;  the  sub-family  Gerbillince,  O.  E.  H.,  is  restricted 
to  Eastern  Arctogaea,  while  the  Murinae  are  exclu- 
sively confined  to  the  Old  World,  but  range  into 
Notogaea. 

Myoxidce.     H.  E. 

Spalacida.     H.  O.  E. 


V.]  SUMMARY   OF   THE   FAUNA.  189 

Hystricidae  ;  in  this  family  the  Hystricina  are  practically 
restricted  to  Eastern  Arctogaea,  although  a  Javan 
species  is  found  in  Timor. 

UNGULATA. 

Hippopotamidce.     E.  O.f  H.f  M.f 

Suidce.  H.  O.  E.  M.,  also  extending  into  the  Austro- 
Malayan  region. 

*  Anoplotheriida.     H. 

*  Dichodontidce.     H. 

Camelidae ;  in  this  group  the  genus  Camelus,  H.  E.  O.,  is 

peculiar  to  Eastern  Arctogaea. 
Tragulidae ;  the  existing  genera  Tragulus,  O.,  and  Dorca- 

therium,  E.  H.t,  as  well  as  several  extinct  ones, 

are  restricted  to  this  area. 
Giraffida.     E.  O.f  H.f 
Bovidae;  the  whole  of  the  true  antelopes  and  goats,  as 

well  as  most  of  the  sheep  and  oxen,  are  restricted  to 

Eastern  Arctogaea,  North  America  now  preserving 

only  one  species  of  Bos,  Ovis,  Ovibos,  and  Haplo- 

ceros. 

*  Palaeotheriidae ;  Palceotherium. 

*  Lophiodontidae ;  Lophiodon. 
Procaviida.     E.  H. 

*  Dinotheriida.     H.  O. 

EFFODIENTIA. 

Manida.     E.  O.  H.f 
Orycteropodida.     E.  H.f 

To  these  may  be  added  the  absence  of  living  opossums  (Didel- 
phyida}.  It  will  be  observed  that  in  this  list  such  existing  families 
as  are  confined  to  one  of  the  regions  of  the  area  considered  are 
for  the  most  part  omitted.  Tertiary  families  which  are  at  present 
unknown  beyond  the  Eastern  Holarctic  region  have,  however,  been 
included,  since  the  limitation  is  probably  in  great  part  due  to  our 
want  of  knowledge  of  the  early  Tertiary  faunas  of  the  other 
regions. 

Although   the    differences   indicated   between   the   faunas   of 


190  EASTERN   ARCTOG^A.  [CHAP. 

Eastern  and  Western  Arctogaea,  which  will  be  still  more  apparent 
after  the  consideration  of  the  mammals  of  North  America,  seem  at 
first  sight  to  indicate  that  these  two  areas  should  form  distinct 
divisions  of  the  realm,  yet  the  community  of  the  fauna  of  the 
northern  portion  of  the  two  hemispheres  forbids  this  view.  This 
question  may,  however,  be  more  fully  discussed  in  the  chapter 
devoted  to  the  Holarctic  region. 

Before  entering  upon  the  consideration  of  the  different  zoolo- 
gical regions  into  which  the  realm  is  divided,  it  is 
MlmmarHan         essential  to  take  a  brief  survey  of  the  Tertiary  mam- 
Faunas  of  malian  faunas  of  Eastern  Arctogaea.     By  this  alone 

Eastern  . 

Arctogeea.  it  is  possible  to  understand  the  true  relations  of  the 

existing  faunas  to  one  another ;  while  such  a  survey 
also  serves  to  demonstrate  that  the  regions  in  question  are  but 
features  of  the  present  epoch  of  the  earth's  history ;  and  that  even 
as  late  as  the  Pliocene  portion  of  the  Tertiary  epoch  the  dis- 
tinctions now  obtaining  between  the  Holarctic,  Oriental,  and 
Ethiopian  regions  had  no  existence.  In  our  survey  we  may  omit 
the  Eocene  period,  and  commence  with  the  lower  Oligocene  ;  and 
it  will  simplify  matters  to  give  lists  of  some  of  the  more  important 
and  better  known  generic  types  characterising  the  faunas  of  the 
different  horizons.  The  leading  affinities  of  many  of  the  genera 
mentioned  have  been  already  alluded  to  in  the  present  or  preced- 
ing chapters  ;  but  it  would  vastly  exceed  the  limits  of  our  space 
to  attempt  to  point  out  the  distinctive  features  of  the  others. 
Accordingly,  the  reader  must  either  take  them  on  trust,  and  treat 
them  practically  as  abstract  terms,  or  he  must  refer  to  some 
palaeontological  treatise  in  order  to  find  the  real  nature  of  the 
animals  indicated  by  such  generic  names. 

Although  it  is  essential  to  our  purpose  to  notice  the  Oligocene 

faunas    of  Eastern    Arctogaea,   it   is    important    to 

FauIS*06116         observe  that  our  knowledge  of  these  is  practically 

limited  to  Western  Europe.    We  are  consequently 

quite   unable   to    say    how   far    the    geographical  range    of  such 

faunas  extended,  although  it  is   probable  that  this   embraced  a 

large  portion  of  the  Eastern  Holarctic  region.    Whether,  however, 

at  this  epoch  Ethiopian  Africa  had  received  a  large  mammalian 

fauna  must  be  left  for  future  discoveries  to  determine. 


V.]  OLIGOCENE   FAUNA.  Ipl 

Under  the  term  of  lower  Oligocene  (the  upper  Eocene  of 
many  writers)  are  included  a  large  series  of  strata,  such  as  the 
freshwater  beds  of  Bembridge  and  Hordwell  in  the  south  of 
England,  the  gypsum  of  Montmartre  near  Paris,  and  the  corre- 
sponding black  lignite  beds  of  De'bruge  in  Vaucluse1.  A  consider- 
able part  of  the  fauna  of  the  Quercy  phosphorites  of  Central  France 
likewise  comes  under  the  same  category,  only  we  have  here  a 
mixture  of  Middle  and  Upper  Oligocene  forms.  And  in  the  case 
of  the  siderolites,  or  bone-earths  of  Switzerland,  this  admixture  is 
carried  to  a  still  greater  degree,  undoubted  Eocene  types  occurring 
with  those  properly  characteristic  of  the  Oligocene.  In  the 
following  list  such  genera  as  are  found  only  in  the  phosphorites 
have  the  letter  P.  after  them;  while  after  those  peculiar  to  the 
siderolites  the  letter  S.  is  added;  both  letters  being  given  when 
the  genera  are  common  to  the  two  formations.  As  already  said, 
only  some  of  the  better-known  forms  are  selected. 

Among  the  lemuroid  Primates,  we  have  the  genera  Adapts 
and  Microch&rus,  both  of  which  occur  in  the  English  beds  as  well 
as  in  the  phosphorites ;  these  being  the  last  European  representa- 
tives of  the  group.  The  Insectivora  include  Necrogymnurus  (P.S.), 
allied  to  the  Malayan  Gymnura,  Amphidozotherium,  together  with 
the  existing  genera  Sorex  and  Talpa.  More  remarkable  is  the 
occurrence  in  the  phosphorites  of  an  insectivore  described  as 
Pseudorhynchocyon,  which  is  believed  to  be  a  member  of  the  family 
of  jumping  shrews  (Macroscelidida),  now  confined  to  the  Ethiopian 
region. 

The  true  Carnivora  are  represented  by  Eusmilus  (P.),  a 
highly  specialised  ally  of  the  sabre-toothed  tigers,  as  well  as  by 
the  more  cat-like  sElurictis,  and  the  generalised  Pseudcelurus. 
In  addition  to  species  of  true  civets,  referred  to  the  living  genus 
Viverra,  the  Viverrida  include  Amphictis  (P.),  Stenoplesictis  (P.), 
and  Palaoprionodon  (P.);  the  two  latter  being  generalised  forms 
closely  connecting  the  family  with  the  Mustelidce,  which  is 
represented  by  Plesictis  (P.).  To  the  Canida  may  be  assigned 
the  genera  Cynodon  (P.S.),  Cephalogale  (P.),  and  Cynodictis, 
together  with  a  species  which  may  be  included  in  Amphicyon 

1  See  table  on  p.  117. 


192  EASTERN   ARCTOG^A.  [CHAP. 

(P.);  while  the  creodont  division  of  the  order  is  represented  by 
Hyanodon,  Pterodon,  Oxycena  (P.),  and  Proviverra.  Among  the 
rodents  we  may  note  the  squirrel-like  Sciuroides  (P.S.)  and  Pseudo- 
sciurus  (S.),  the  existing  genus  Sa'urus,  and  the  extinct  Plesiarctomys 
and  Plesiospermophilus,  which  likewise  belong  to  the  same  family. 
Among  the  Muridce,  Cricetus  includes  ancestral  types  of  the 
hamsters ;  while  the  dormice  are  represented  by  the  existing  genus 
Myoxus.,  As  noticed  previously,  Theridomys,  Nesocerodon,  and 
Protechinomys  seem  to  be  ancestral  forms  allied  to  some  of  the 
existing  South  American  rodents. 

The  ungulates  are  very  strongly  represented;  the  pig-like 
group  including  Cebochoerus,  Cheer opotamus  and  Elotherium  (P.) 
among  the  Chcsropotamida^  Acotherulum,  and  Anthracotherium  (P.) 
and  Ancodus  in  the  Anthracotheriidce.  The  anoplotheroids  com- 
prise Anoplotherium,  D aery t her ium,  and  Xiphodon  ;  Dichobunus 
and  Ccenotherium  (P.)  are  the  characteristic  forms  among  the 
Ccenotheriidce;  and  Dichodon,  Gelocus,  and  Lophiomeryx  among  the 
DichodontidcR\  while  the  chevrotains  are  represented  by  Prodremo- 
therium  (P.)  and  Bachitherium  (P.).  In  the  perissodactyle  section 
of  the  same  order,  we  have  Pachynolophus  (P.)  to  represent  the 
Lophiodontidce,  Palceotherium  and  Anchilophus  in  the  Palceo- 
theriida,  Protapirus  (P.)  as  an  ancestral  form  of  the  tapirs,  and 
Rhinoceros  (P.),  Cadurcotherium  (P.),  and  Hyrachyus  (P.)  as 
representatives  of  the  rhinoceroses.  The  aberrant  Chalicotherium 
has  also  one  species  from  the  phosphorites.  The  Effodientia  in- 
clude Leptomanis  (P.),  Necromanis  (P.),  and  Palceorycteropus  (P.); 
while  the  existing  genus  Didelphys  alone  represents  the  mar- 
supials. 

It  will  be  seen  that  in  this  fauna  the  existing  generic  types  are 
very  few,  and  if  the  whole  of  the  extinct  ones  had  been  given,  their 
relative  proportion  would  have  been  still  less.  The  ungulates  were 
abundant,  and  among  these  the  perissodactyles  proportionately 
more  numerous  than  at  the  present  day ;  while  the  anoplotheres 
are  in  some  respect  transitional  between  the  latter  and  the  typi- 
cal artiodactyles.  All  the  ungulates  had  brachydont  teeth,  and 
annectant  types  between  the  modern  pigs  and  ruminants  were 
abundant ;  the  traguloids  being  the  highest  development  among 
the  artiodactyle  section  of  the  order.  Creodont  carnivores  still 


V.]  OLIGOCENE   FAUNA.  193 

persisted,  although  more  modern  types  had  already  made  their 
appearance  on  the  scene ;  and  opossums  flourished. 

The  middle  stage  of  the  Oligocene  is  represented  in  Europe 
by  the  freshwater  marls  and  clays  of  Hempsted  in  the  Isle  of 
Wight  and  the  corresponding  beds  of  Ronzon,  near  Puy-en-Velay, 
the  lignitiferous  strata  of  Cadibona  in  Liguria,  the  deposits  of 
Fontainebleau  and  Ferte-Alais  in  France,  and  likewise  by  certain 
beds  in  Hungary  and  at  Monte  Promina  in  Dalmatia.  Among 
the  small  fauna  of  this  stage  we  may  notice  the  following.  In  the 
Insectivora,  Tetracus,  an  ally  of  the  hedgehogs;  Cynodon,  Amphi- 
cynodon,  Plesictis,  and  Hycznodon  among  the  carnivores;  Anthra- 
cotherium,  Ancodus,  Elotherium,  Ccenotherium,  Gelocus,  and  Rhino- 
ceros in  the  ungulates;  and  opossums  (Didelphys).  While  this 
fauna  is  closely  related  to  the  preceding,  it  has  lost  a  number  of 
early  ungulate  types,  such  as  Anoplotherium  and  Xiphodon  among 
the  artiodactyles,  and  Pal&otherium  and  Anchilophus  in  the 
perissodactyles.  On  the  other  hand,  the  pig-like  forms,  such  as 
Ancodus,  Anthracotherium,  and  Elotherium,  attained  an  extra- 
ordinary degree  of  development.  Among  the  creodont  carnivores, 
we  may  note  the  final  disappearance  of  the  genera  Pterodon  and 
Proviverra,  although  Hycenodon  still  survived. 

The  upper  Oligocene  (lower  Miocene)  fauna  is  a  large  and 
characteristic  one,  well  represented  in  the  freshwater  beds  of 
St  Gerand  le  Puy,  in  the  Allier,  as  well  as  in  those  of  Weisenau 
and  other  localities  in  the  neighbourhood  of  Mayence.  Among 
the  mammals  may  be  mentioned  the  following ;  existing  genera 
being  denoted  by  the  prefix  of  a  f. 

INSECTIVORA.  fTalpa.  fSorex. 

Geotrypus.  Dimylus. 

t  Myogale.  Palseoerinaceus. 

Plesiosorex.  t  Erinaceus. 

CARNIVORA.  Cephalogale.  fViverra. 

Amphicyon.  t  Herpestes. 
Plesictis.  Proaelurus. 

Potamotherium.  Hyaenodon. 

Amphictis. 

L.  13 


194  EASTERN   ARCTOGyEA.  [CHAP. 

RODENTIA.  Theridomys.  fSpermophilus. 

Archaeomys.  f  Sciurus. 

Issiodoromys.  Chalicomys. 

t  Myoxus.  Titanomys. 

t  Cricetus. 

UNGULATA.  Anthracotherium.         Csenotherium. 

Hyotherium.  f  Tapirus. 

Amphitragulus.  t  Rhinoceros. 

Dremotherium. 
MARSUPIALIA.          f  Didelphys. 

Of  this  fauna,  Professor  von  Zittel  writes  that  it  seems  at  first 
sight  closely  akin  to  those  of  the  middle  and  lower  Oligocene ;  the 
same  ordinal  and  subordinal  groups,  and  in  many  instances  the 
same  genera  characterising  the  whole  three  horizons.  In  the  lack 
of  lemuroids,  the  reduced  number  or  final  disappearance  of 
opossums,  creodonts,  and  anoplotherioids,  in  the  greater  abundance 
of  forms  like  Anthracotherium,  Hyotherium,  and  Dremotherium, 
which  were  but  poorly  represented  in  the  lower  Oligocene,  and  in 
the  number  of  new  types,  such  as  Tapirus,  Amphitragulus  (an 
ancestral  chevrotain),  Chalicomys  (an  early  beaver),  Titanomys  (an 
ally  of  the  picas),  Erinaceus,  Dimylus  (a  form  connecting  the 
shrews  with  the  hedgehogs),  Potamotherium  (a  generalised  otter), 
Herpestes  (mungooses),  Procelurus  (a  primitive  type  of  cat),  we 
notice,  however,  the  marked  difference  of  this  fauna  from  its  fore- 
runners. Among  the  incoming  genera  it  is  noteworthy  that  there 
is  none  for  which  an  ancestral  type  cannot  be  found  in  the  lower 
Oligocene ;  the  main  difference  occurring  in  the  more  specialised 
characters  of  the  members  of  the  later  fauna.  With  the  exception 
of  certain  bats,  insectivores,  rodents,  and  the  opossums  (such  as 
Vespertilio,  Erinaceus,  Sorex,  Myogale,  Talpa,  Sciurus,  Spermo- 
philus,  Cricetus,  Myoxus,  and  Didelphys],  the  majority  of  the 
genera  are,  however,  still  extinct. 

It  is  probable  that  the  beds  in  the  Balkans  which  have  yielded 
remains  of  the  North  American  Tertiary  genus  Titanotherium 
belong  to  some  portion  of  the  Oligocene  epoch. 

We  now  come  to  the  Miocene  epoch,  which,  as  at  present  re- 
stricted, forms  in  Europe  but  a  small  section  of  the  Tertiary  era. 


V.]  MIOCENE    FAUNA.  195 

It  includes  the  well-known  freshwater  strata  of  Sansan  in  Gers  (the 
middle  Miocene  of  the  older  geological  classifica- 
tions),   together   with    the   corresponding   beds    of     F^n°acene 
Steinheim    in    Styria,   and   likewise    the   somewhat 
newer  (upper  Miocene)  deposits  of  CEningen,  in  Baden.    Grive-St- 
Alban,  in  the  valley  of  the  Rhone,  is  likewise  another  well-known 
locality  where  mammaliferous  strata  of  this  age  are  developed; 
and,  among  other  places,  we  may  also  mention  Monte  Bamboli 
in  Italy,  San  Isidro  in  Spain,  and  Oran  in  Algeria. 

For  the  first  time  in  Europe  we  meet  with  remains  of  true 
Primates,  of  which  there  are  three  genera  belonging  to  the  Simiidce, 
two  of  which,  Dryopithecus  and  Oreopithecus *,  are  extinct,  but  the 
third  seems  scarcely  separable  from  the  existing  Oriental  Hylobates. 
In  the  Insectivora  we  meet  with  the  existing  European  genera 
Talpa,  Myogale  (desmans),  Erinaceus,  Soreoc,  and  Crocidura ;  while 
the  extinct  Lanth  another  turn  seems  to  be  allied  to  the  tree-shrews 
( Tupaid]  of  the  Oriental  region,  and  Galerix  intermediate  between 
the  latter  and  the  j Limping-shrews  (Macroscelidtdce)  of  Ethiopian 
Africa.  Among  the  Carnivora,  where  the  creodonts  have  disap- 
peared, the  cats  are  represented  by  the  sabre-toothed  tigers 
(Machcerodus]  and  Pseudcelurus.  In  addition  to  the  existing 
genera  Viverra  and  Herpestes,  we  have  among  the  civet  tribe  the 
extinct  Progemtta.  The  dogs  include  the  existing  Cam's,  together 
with  the  extinct  Hemicyon  and  Pseudocyon ;  while  the  larger  forms 
described  as  Dinocyon  and  Hycznarctus  connect  the  former  with 
the  bears.  The  Mustelidce.  are  represented  by  species  of  the 
typical  living  genus  Mustela,  together  with  certain  more  or  less 
closely  allied  extinct  types ;  and  Enhydriodon  filled  the  place 
of  the  modern  otters. 

From  among  the  rodents  the  generalised  types  allied  to  those 
now  characteristic  of  Neogsea  have  all  disappeared,  nearly  all  the 
recorded  forms  apparently  pertaining  to  existing  genera.  In  the 
Sciuridce  not  only  have  we  true  squirrels  (Sdurus\  but  the  Ethio- 
pian spiny  squirrels  (Xerus)  are  likewise  represented,  as  are  also 
the  more  widely  distributed  flying-squirrels  of  the  genus  Stiuro- 
pterus,  which  now  inhabit  both  Eastern  and  Western  Arctogaea 

1  Some  of  the  characters  of  these  genera  have  been  already  mentioned  on 
pages  1 80,  18 r. 

13—2 


196  EASTERN    ARCTOG^A.  [CHAP. 

Chalicomys,  Cricetus,  and  Myoxus  are  survivors  from  the  Oligocene; 
but  porcupines  (Hystrix]  are  new  comers.  Picas  of  the  existing 
genus  Lagomys  are  likewise  to  the  fore ;  and  it  is  a  question 
whether  those  distinguished  by  the  name  of  Myolagus  might  not 
be  included  under  the  same  title. 

A  marked  approximation  to  the  modern  type  is  likewise  the 
characteristic  feature  of  the  ungulates  of  the  European  Miocene ; 
although  in  this  group  living  genera  still  remain  in  the  minority. 
The  pigs  (Sutda)  include,  for  instance,  the  genus  Hyotherium,  in 
which  the  molar  teeth  are  tuberculated  and  of  the  general  type  of 
those  of  some  of  the  living  members  of  the  family ;  and  also  the 
more  aberrant  Listriodon,  characterised  by  the  presence  of  a  pair 
of  transverse  ridges  on  each  of  the  teeth  of  the  same  series.  A 
species  of  the  existing  West  African  genus  Dorcatherium  alone 
represents  the  chevrotains  (Tragulidte) ;  while  we  have  forerunners 
of  the  deer  (Cervidce)  in  the  extinct  Pal&omeryx  and  Dicroceros, 
both  characterised  by  the  simple  structure  of  their  antlers ;  and 
Protragoceros — a  generalised  type  of  antelope — marks  the  first 
appearance  of  the  hollow-horned  ruminants  (Bovida],  which  now 
form  such  a  numerous  and  characteristic  group  in  the  fauna  of 
Eastern  Arctogaea.  Perissodactyle  ungulates  are  less  numerous. 
Anchitheriiim,  to  some  of  whose  distinctive  characters  allusion  has 
been  made  above,  constitutes  the  representative  of  the  equine  line 
at  this  stage ;  and  tapirs  and  rhinoceroses  belonging  to  the 
existing  genera  were  likewise  common.  Some  of  the  latter  were, 
however,  still  hornless,  and  in  none  was  more  than  a  single  horn 
developed.  The  aberrant  ChalicotheriidcB^  forming  the  last  family 
of  this  section  of  the  order,  and  characterised  by  the  extraordinary 
resemblance  presented  by  their  claws  and  toes  to  those  of  eden- 
tates, are  here  represented  by  the  gigantic  Macrotherium.  Finally 
the  Miocene  is  notable  as  being  the  stage  at  which  proboscideans 
first  made  their  appearance  on  the  scene  in  Europe.  In  this  group 
we  have  species  of  Mastodon,  which,  as  already  explained,  includes 
the  ancestors  of  the  modern  elephants ;  and  likewise  one  of  the 
more  aberrant  Dinotherium. 

Compared  with  the  Oligocene,  the  loss  of  so  many  antiquated 
types,  coupled  with  the  appearance  of  proboscideans  and  man-like 
apes,  and  the  general  modern  facies  of  all  the  mammals  of  the: 


V.]  LOWER   PLIOCENE   FAUNAS.  197 

Miocene,  indicates  the  lapse  of  a  considerable  interval  of  time 
between  the  deposition  of  the  two  series  of  strata.  And  that  this 
is  really  the  case,  is  demonstrated  by  the  fact  that  there  occurs 
between  the  two  a  considerable  thickness  of  marine  deposits  which 
have  not  hitherto  yielded  remains  of  land  mammals.  It  may  be 
noticed  that  while  many  of  the  insectivores  and  rodents  from  this 
horizon  belong  to  genera  now  inhabiting  the  Eastern  Holarctic 
region,  among  other  forms  we  have  marked  instances  of  Oriental 
(Lanihanotherium.  Hylobates]  or  Ethiopian  ( Galerix,  Dorcatherium, 
and  Xerus)  affinities  in  this  assemblage;  and  it  is  thus  evident  that 
at  the  epoch  in  question  there  was  no  trace  of  the  differentiation 
of  Eastern  Arctogaea  into  regions. 

Still  more  markedly  are  the  same  features  displayed  by  the 
older  Pliocene  fauna  of  Europe  and  Southern  Asia. 
This  fauna,  which  was  formerly  regarded  as  of  upper  cen^Faun'a 
Miocene  age  until  shewn  by  Dr  Blanford  to  be 
unquestionably  referable  to  the  succeeding  era  of  geological  his- 
tory, had  a  very  wide  distribution  ;  and  it  is  represented  at  certain 
localities,  mostly  at  long  distances  from  one  another,  by  an  extra- 
ordinary profusion  of  remains.  One  of  these  charnel-holes  occurs 
at  the  village  of  Pikermi,  near  Athens,  a  second  in  the  Isle  of 
Samos,  in  the  Turkish  Archipelago,  and  a  third  at  Mont  Leberon, 
in  Provence.  This  fauna  is  also  met  with  locally  in  the  valley  of  the 
Rhone,  at  the  foot  of  the  Pyrenees,  in  Spain,  Asia  Minor,  and  at 
Maraga  in  Persia.  It  is  likewise  represented  in  the  regions  lying 
to  the  north  of  the  Alps,  only  here  the  number  of  forms  is  less, 
and  the  antelopes  and  giraffe-like  ruminants,  fitted  for  roaming 
over  the  open  plains  of  the  south,  are  conspicuous  for  their 
absence;  their  place  being  taken  by  forest-haunting  deer.  The 
sand-beds  of  Eppelsheim  in  Hessen-Darmstadt,  together  with 
strata  in  the  neighbourhood  of  Vienna,  and  others  in  Hungary 
and  Rumania,  may  be  cited  as  localities  where  the  northern 
section  of  this  fauna  is  preserved. 

Taking  first  the  European  and  Western  Asiatic  portion  of  this 
fauna,  and  leaving  its  Oriental  members  for  subse- 
quent consideration,  we  find  the  Primates  represented 
solely  by  an   extinct  genus  of  monkey,  taking  its 
name   of  Mesopithecus  on  account  of  presenting  certain  features 


198  EASTERN   ARCTOG^A.  [CHAP. 


intermediate  between  the  existing  Semnopithecus  and  Macacus. 
The  insectivores  are  likewise  known  only  by  a  solitary  form,  a 
shrew  (Sorex) ;  but  this  is  probably  due  to  the  nature  of  the  strata 
being  unfitted  for  the  preservation  of  the  remains  of  such  small 
creatures.  The  Carnivora,  on  the  other  hand,  were  abundant,  the 
Felidce  being  represented  not  only  by  the  sabre-tooths  (Machter- 
odus),  but  true  cats  (Felis)  likewise  making  their  appearance  on 
the  scene.  Hyaenas  display  a  great  variety  of  development,  there 
being  one  species  of  the  typical  genus  Ifycena,  with  certain 
resemblances  to  the  existing  Cape  form,  while  the  more  generalised 
types  known  as  Lycyana  and  Hycenictis  were  likewise  present,  as 
were  also  species  of  Ictitherium^  and  the  allied  Palhyana,  which, 
as  already  mentioned,  formed  a  connecting  link  between  the 
hyaenas  and  the  civets.  True  dogs  seem  to  have  been  absent  from 
this  assemblage ;  but  Amphicyon  still  survived  from  the  Miocene, 
and  an  aberrant  form  known  as  Simocyon  made  its  appearance. 
Hycznarctus  was  likewise  another  survivor  from  the  Miocene,  and 
may  be  regarded  as  a  forerunner  of  the  true  bears.  Finally,  in  the 
weasel  tribe  (MustelidcB)  we  have  representatives  of  the  existing 
genus  Mustela,  as  well  as  the  extinct  Pal&omephitis  and  Promeles, 
the  latter  being  an  ancestral  type  of  the  badgers. 

Rodents  make  but  a  poor  show,  as  we  have  only  the  extinct 
beaver-like  Chalicomys,  a  species  of  porcupine  (ffystrix),  and  a 
representative  of  the  curious  little  spiny  mice  (Acomys\  now 
characteristic  of  Syria,  Palestine,  and  north-eastern  Africa. 

A  remarkable  advance  over  their  Miocene  forerunners  is 
displayed  by  the  ungulates,  especially  those  from  Pikermi  and 
Maraga.  Here,  in  the  artiodactyle  section,  we  meet  for  the  first 
time  with  true  pigs  of  the  genus  Sus,  which  at  this  period  ranged 
over  the  greater  portion  of  Europe,  and  some  of  which  attained 
very  large  dimensions.  Water-chevrotains  (Dorcatherium)  serve 
to  connect  the  Miocene  representatives  of  their  genus  with  the 
existing  West  African  form;  while  muntjacs  (Cervulus),  now 
confined  to  the  Oriental  region,  filled  the  place  of  the  stags. 
Giraffe-like  creatures  were  numerous,  for  not  only  have  we  true 
giraffes  belonging  to  the  existing  Ethiopian  genus  Giraffa,  but  the 
gigantic  hornless  Helladotherium  stalked  over  the  plains  of  Greece, 
and  the  allied  but  horned  Samotherium  inhabited  the  area  now 


LOWER   PLIOCENE    FAUNAS. 


199 


occupied  by  the  Turkish  Archipelago,  and  extended  eastwards  as  far 
as  Persia;  Palceotragus  being  a  smaller  but  allied  form.  The  Bovidce 
are  represented  by  antelopes,  most  of  which  present  a  marked 
Ethiopian  facies,  although  Tragoceros  (probably  the  direct  descen- 
dant of  the  Miocene  Protragoceros}  is  an  aberrant  form,  with 
compressed  horn-cores  like  those  of  the  goats.  And  it  may  be 
remarked  that  most  of  the  Pikermi  antelopes  have  short-crowned 
molar  teeth,  in  which  respect  they  resemble  the  existing  eland, 
kudu,  and  their  allies.  Of  the  Pliocene. forms,  Palczorias,  which  is 
common  to  Southern  Europe  and  Algeria,  seems  to  be  inter- 


FlG.   45.    SKULL  OF  Palczorias. 

mediate  between  the  kudus  (Strepsiceros)  and  elands  (Orias}\ 
while  the  so-called  Protragelaphus  is  so  closely  allied  to  the 
existing  Ethiopian  harnessed  antelopes  (Tragelaphus)  as  to  be 
included  by  some  in  the  same  genus.  On  the  other  hand, 
Palaoryx  is  nearly  related  to  the  gemsbok  and  its  allies  (Oryx), 
although  with  certain  resemblances  to  the  sable  antelope  group 
(Hippotragus).  Gazelles  (Gazella},  which  are  essentially  inhabi- 


2OO  EASTERN   ARCTOG^A.  [CHAP. 

tants  of  open  plains,  were  likewise  abundant ;  one  being  considered 
a  near  relative  of  the  South  African  springbok.  The  genus 
Helicophora,  on  the  other  hand,  closely  resembles  the  water-buck 
group  (Cobus),  which  is  exclusively  Ethiopian.  In  the  perisso- 
dactyle  division,  the  three-toed  horses  (Hipparion)  seem  to  have 
approximated  in  general  structure  to  the  Ethiopian  zebras,  and, 
like  those  animals,  may  have  been  ornamented  with  dark  and  light 
stripes.  While  some  of  the  Pliocene  rhinoceroses  were  hornless, 
another  was  a  two-horned  species  closely  allied  to  the  common 
African  Rhinoceros  bicornis,  of  which  it  may  be  regarded  as  the 
parent  form.  There  is  also  an  extinct  genus  (Leptodon),  of  some- 
what uncertain  affinity ;  while  tapirs  are  found  in  the  Eppelsheim 
beds,  although  not  apparently  in  the  southern  area.  The  Chalico- 
theriidft  were  represented  by  the  typical  genus  Chalicotherium 
(Ancylotheriuni),  which,  as  we  have  seen,  was  a  near  ally  of  the 
Miocene  Ma  cr other  him,  and  also  occurs  in  the  Oligocene  phos- 
phorites. As  in  the  Miocene,  the  proboscideans  include  only 
Mastodon  and  Dinotherium;  the  one  species  of  the  former  ranging 
from  Greece  to  Persia,  but  being  different  from  all  the  Indian  forms 
of  the  same  epoch.  Finally,  the  occurrence  of  an  aard-vark 
(Orycteropus]  both  in  Samos  and  Persia  serves  to  accentuate  the 
Ethiopian  affinities  of  the  southern  section  of  this  fauna. 

We  have  thus  evidence  that  one  and  the  same  fauna  extended 
from  Spain  and  Algeria  across  Southern  Europe  to  Asia  Minor 
and  Persia;  and  we  may  infer  from  the  deposits  at  Samos,  that 
what  is  now  the  y-Egean  sea  formed  a  tract  of  land  connecting 
Greece  with  Turkey.  It  is  further  evident  that  there  must  have 
been  free  communication  across  the  Mediterranean  basin  (which 
in  Cretaceous  times  is  known  to  have  been  a  mare  clausnm,  in  the 
physical,  and  not  the  political  sense  of  the  term)  between  Europe 
and  Africa.  This  communication  may  have  existed  both  by  way 
of  Gibraltar,  and  also  between  Italy,  Sicily,  and  Malta  on  the 
one  hand,  and  Tunis  on  the  other;  since  the  Plistocene  mammals 
of  the  islands  in  question  clearly  indicate  continental  connection. 
While  the  antelopes  and  hipparions  of  this  fauna  prove  the  exist- 
ence of  open  plains  during  the  lower  Pliocene  epoch,  the  host  of 
individuals  of  Mesopithecus  as  unmistakeably  point  to  the  presence 
of  extensive  forest-tracts.  In  the  northern  section  of  the  fauna,  as 


V.]  SIWALIK   FAUNA.  2OI 

displayed  at  Eppelsheim,  the  Ethiopian  affinities  are  much  less 
apparent,  aard-varks  and  the  whole  of  the  giraffe-group  being 
absent,  while  tapirs  and  deer  were  abundant.  That  there  was  a 
more  or  less  marked  separation  between  the  two  areas  thus  seems 
evident;  and  the  tapirs  and  muntjac-like  deer,  both  of  which 
seem  wanting  in  the  Siwalik  fauna,  are  indicative,  so  far  as  they 
go,  of  Malayan  affinities. 

Nearly  related  to   that    of   Pikermi,   Samos,   and  Persia,  the 
celebrated  Siwalik  fauna  of  India  and  the  adjacent 
countries  presents   certain  well-marked  differences;     Fa^hk 
this  being  specially  shown  by  the  occurrence  of  several 
essentially  modern   types  quite  unknown  in  the  former.     More- 
over, there  are  a  considerable  number  of  peculiar  genera  which  do 
not  occur  in  the  western  fauna;  while  we  also  come  across  certain 
Miocene,  and  even  Oligocene  types,  which  are  equally  strange  to 
the  latter.     Although  in  some  cases  these  occur  in  beds  which  are 
not  improbably  of  upper   Miocene  age,   in   others  they  appear 
mingled  with  the  later  forms ;  but,  in  any  case,  they  indicate  a 
survival  in  this  area  of  archaic  types  which  at  that  time  had  com- 
pletely disappeared  from  Europe. 

Originally  discovered  in  the  outer  ranges  of  the  typical  Hima- 
layan area,  the  Siwalik  fauna  has  been  traced  towards  the 
north-west  into  the  Punjab,  Kach,  Sind,  and  the  north-eastern 
frontier  of  Baluchistan  ;  the  beds  from  the  two  latter  areas  being 
lower  in  the  series  than  those  from  the  typical  Siwalik  hills,  and 
containing  an  older  assemblage  of  forms,  although  several  are 
common  to  all.  An  outlier  of  the  same  fauna  occurs  in  Perim 
Island  in  the  gulf  of  Cambay.  Eastwards  the  Siwalik  fauna 
ranged  through  Sylhet  and  Assam  to  Burma,  whence  it  has  been 
traced  at  intervals,  as  in  Java,  Sumatra,  and  the  Philippines,  into 
China  and  Japan.  In  China  it  extended  from  Yunnan  in  the 
south-west  northwards  through  Szechuen  to  Kansu,  and  thence 
eastwards  through  Shensi  to  Shansi,  its  extreme  eastern  limit  being 
indicated  by  the  discovery  of  a  Siwalik  elephant's  tooth  at 
Shanghai.  Northward  of  Kansu  the  fauna  ranged  into  Mongolia, 
probably  by  way  of  the  gap  formed  by  the  course  of  the  Hwang-ho 
through  the  Ala-shan  mountains — if  such  mountains  existed  at  the 
time.  And  it  is  not  a  little  remarkable  that  of  the  few  Mongolian 


202  EASTERN    ARCTOG^A.  [CHAP. 

forms  at  present  known,  two  (Hycena  macrostoma  and  Equus 
sivalensis)  are  identical  with  species  from  the  Siwalik  Hills1. 

As  regards  the  fauna  itself,  we  find,  in  the  first  place,  the 
Primates  much  more  fully  represented  than  at  Pikermi,  and  all  by 
existing  generic  types.  Of  the  man-like  apes  (Simiid&\  there  is 
a  chimpanzee  (Anthropopithecus)  presenting  a  more  human  type 
of  dentition  than  its  living  Ethiopian  cousins;  while  a  single  tusk 
indicates  the  former  existence  of  an  orang  (Simla)  allied  to  the  living 
Bornean  and  Sumatran  species.  The  other  three  generic  types 
belong  to  the  Cercopithecida,  and  include  baboons  of  the  Ethiopian 
genus  Papio  ( Cynocephalus),  together  with  species  of  Semnopithecus 
and  Macacus,  the  former  genus  being  exclusively,  and  the  latter 
mainly,  Oriental  at  the  present  day,  although  both  occur  in  the 
later  Pliocene  of  Europe.  Doubtless  owing  to  the  unsuitability 
of  the  strata  for  the  preservation  of  small  specimens,  no  remains 
of  insectivores  have  hitherto  been  obtained.  The  Carnivora  are, 
on  the  other  hand,  well  represented;  the  Felicia  including  large 
and  small  species  of  the  typical  genus  Felis,  and  apparently  one 
of  the  allied  Cynahirus  (hunting-leopard),  now  exclusively  Oriental 
and  Ethiopian.  Mach&rodus  had  two  species ;  and  another 
form  has  been  identified  with  the  European  Oligocene  genus 
SEhiridis.  Civets  include  species  of  Viverra  larger  than  any  now 
existing;  this  genus  being  also  one  now  confined  to  the  Ethiopian 
and  Oriental  regions,  although  more  abundant  in  the  latter  than  in 
the  former.  The  CanidcE,  in  addition  to  a  survivor  of  the  Miocene 
Amphicyon,  were  represented  by  wolves  and  jackals  (Cam's),  as 
well  as  by  a  species  apparently  allied  to  the  long-eared  fox  (Otocyon} 
of  Africa.  While  in  the  Ursidce  the  generalised  Hycenarctus  still 
survived,  true  bears  (Ursus)  make  their  appearance  for  the  first 
time,  the  single  known  Siwalik  species  presenting,  however,  a 
marked  approximation  in  the  characters  of  its  skull  and  dentition 
to  the  Indian  sloth-bear  (Melursus).  Among  the  few  known 
representatives  of  the  Mustelida,  we  have  a  large  marten  (Mustela), 
probably  allied  to  the  living  yellow-throated  Indian  species ;  a 
ratel,  belonging  to  a  genus  (Mellivora)  now  restricted  to  India 
and  Africa;  and  likewise  an  otter  (Lutra)  whose  nearest  affinities 
are  with  an  existing  Sumatran  species.  The  same  family  also 

1  Lydekker,  Rec,  Geol.  Sw~v.  India,  Vol.  xxiv.  pp.  207 — 211  (1891). 


V.]  SIWALIK   FAUNA.  203 

includes  a  member  of  the  otter-like  genus  Enhydriodon,  the  other 
species  being  from  the  Italian  Miocene.  Among  the  most 
remarkable  features  of  the  Siwalik  Carnivora  is  the  survival  of 
a  species  of  Hycenodon,  of  which  the  remains  have  been  discovered 
in  the  Punjab. 

The  Rodentia  are  but  very  imperfectly  known.  They  include 
a  representative  of  the  bamboo-rats  (Rhizomys),  which  are  now 
exclusively  Oriental,  and  belong  to  the  family  Spalacida;  and, 
among  the  Muridce,  a  species  of  Nesocia,  which  genus  is  likewise 
confined  to  the  Oriental  region.  The  other  forms  are  a  porcupine 
(Hystrix]  and  a  hare  (Lepus). 

A  very  long  list  is  presented  by  the  ungulates,  which  are 
numerous  not  only  in  generic,  but  likewise  in  specific  types.  The 
pig-like  artiodactyles  include,  among  the  family  Suidce.  several 
representatives  of  the  true  pigs  (Sus),  some  of  which  attained 
gigantic  dimensions,  while  others  are  remarkable  for  the  complex 
structure  of  their  molar  teeth,  which  show  a  marked  resemblance 
to  those  of  the  existing  Ethiopian  wart-hogs  (Phacochczrus).  A 
still  more  elaborate  structure  is  displayed  by  the  corresponding 
teeth  of  the  allied  genus  Hippohyus,  which  is  peculiar  to  this 
fauna ;  and  the  family  is  also  represented  by  species  of  the 
European  Miocene  genera  Hyotherium  and  Listriodon,  the  remains 
of  the  two  latter  being  mostly  obtained  from  the  Punjab  and 
districts  to  the  west.  The  same  areas  are  mainly  those  which 
have  yielded  remains  of  Anthracotheriidcz,  although  some  of  these 
have  been  discovered  in  Sylhet.  In  this  family  we  have  species  of 
the  European  genera  Anthracotherium  and  Ancodus,  the  former  of 


FlG.  46.      RIGHT    UPPER    MOLAR   OF   A   SMALL   SPECIES    OF   Merycopotamns. 

which  is  elsewhere  unknown  above  the  Middle  Oligocene;  and 
there  are  also  three  peculiar  types,  respectively  known  as  Meryco- 
potamus,  Hemimeryx,  and  Chctromeryx,  differing  from  all  the  rest 


204  EASTERN    ARCTOG^A.  [CHAP. 


in  having  only  four  columns  on  the  crowns  of  the  molars,  as 
shown  in  the  annexed  figure,  and  thus  presenting  a  marked 
approximation  to  the  ruminants.  The  earlier  Tertiary  Chczropo- 
tamidcz  likewise  had  a  survivor  in  the  genus  Tetraconodon,  which 
was  represented  by  a  large  pig-like  creature  remarkable  for  the 
enormous  size  of  its  simple  conical  premolar  teeth.  The  pig- 
like  group  closes  with  Hippopotamus,  which  makes  its  appearance 
on  the  scene  for  the  first  time  in  this  formation,  where  it  is 
represented  by  a  generalised  species  with  three  pairs  of  incisor 
teeth  in  each  jaw.  Turning  to  the  groups  with  fully-developed 
selenodont  molars,  we  have  first  to  notice  the  occurrence  of  fossil 
camels  of  the  existing  genus  Camelus,  which  are  unknown  else- 
where except  in  the  Algerian  Plistocene.  As  we  have  seen  that 
the  Camelidce.  were  originally  a  New  World  group,  it  is  interesting 
to  note  that  these  earliest  Old  World  representatives  occur  in  Asia 
instead  of  Europe;  and  it  is  further  noteworthy  that  in  the 
structure  of  their  molar  teeth  the  Siwalik  camels  retain  evidences  of 
affinity  with  the  South  American  guanacos  and  vicunas  which  are 
lost  in  their  living  descendants.  The  Tragulidce  contain  repre- 
sentatives of  the  true  chevrotains  ( Tragulus)  and  water-chevrotains 
(Dorcatherium\  now  respectively  characteristic  of  the  Oriental  and 
Ethiopian  regions,  while  among  the  deer  ( Cervidce)  we  have  species 
of  the  Oligocene  European  genus  Palaomeryx,  together  with 
others  belonging  to  Cervus,  the  representatives  of  the  latter  being 
all  closely  allied  to  existing  Oriental  types.  Not  improbably  also 
a  musk-deer  (Moschus)  should  be  included  among  the  Siwalik 
Cervidce.  Among  the  Giraffidce,  in  addition  to  true  giraffes 
(Giro/a),  which  are  common  to  the  Pikermi  beds,  and  extended 
eastwards  into  China,  we  have  the  peculiar  gigantic  antlered  types  re- 
spectively known  as  Vishnutherium,  Sivatherium,  Hydaspotherium, 
and  Bramatherium,  of  which  the  first  seems  common  to  the  Siwaliks 
of  Burma  and  the  Punjab,  while  the  second  is  confined  to  the 
more  easterly  Himalaya,  the  third  to  the  Punjab,  and  the  fourth  to 
Perim  Island.  They  include  the  most  gigantic  of  all  ruminants, 
Sivatherium  almost  rivalling  an  elephant  in  bulk. 

Not  one  of  the  least  curious  features  in  this  marvellous  fauna  is 
that  while,  as  we  have  seen,  deer  of  Oriental  types  were  abundant, 
antelopes  closely  allied  to  those  now  inhabiting  the  Ethiopian 


v.] 


SIWALIK    FAUNA. 


205 


206  EASTERN    ARCTOG^EA.  [CHAP. 

region — where  deer  are  totally  absent — were  likewise  extraordinarily 
numerous.  Of  the  African  genera  we  have  a  species  of  Bubalis 
intermediate  between  the  hartebeests  and  the  blesbok,  a  member 
of  the  sable  antelope  group  (Hippotragus\  a  kudu  (Strepsiceros),  an 
eland  (Orias),  and  probably  a  representative  of  the  water-buck 
group  (Cobus).  On  the  other  hand,  Oriental  forms  are  not 
wanting,  as  proved  by  the  occurrence  of  a  nilgai  (Boselaphus),  and 
probably  of  a  four-horned  antelope  (Tetraceros] ;  while  the  widely- 
spread  gazelles  (Gazelld)  were  likewise  present.  Goats  and  oxen 
for  the  first  time  made  their  appearance ;  the  former  group  being 
represented  not  only  by  species  belonging  to  the  typical  Capra, 
but  likewise  to  the  shorter-horned  genus  Hemitragus,  now  confined 
to  India  and  Arabia.  The  oxen  (Bos)  included  members  of  all  the 
existing  groups,  that  is  to  say  typical  oxen,  bison,  buffalo,  and 
smaller  forms  with  upright  triangular  horns  nearly  allied  to  the 
anoa  of  Celebes. 

The  perissodactyle  ungulates,  so  numerous  in  the  earlier 
Tertiary  formations,  have  now  become  proportionately  much  fewer 
as  compared  with  the  artiodactyles.  While  typical  forms  of 
Hipparion  were  present,  one  species  differs  from  the  rest  by  the 
loss  of  the  lateral  toes,  and  thus  resembles  the  modern  horses 
(Equus\  which  here  make  their  appearance  for  the  first  time. 
Rhinoceroses  include  not  only  hornless  forms,  but  likewise  one 
species  allied  to  the  existing  Oriental  Rhinoceros  unicornis  and  R. 
sondaicus,  and  a  third  as  closely  related  to  the  African  Burchell's 
rhinoceros  (7?.  simus].  In  the  same  group  Chalicotherium  is  a 
survivor  from  older  formations. 

Finally,  the  proboscideans  exhibit  a  development  unparalleled 
in  any  other  formation  or  epoch.  Dinotherium  appears  for  the 
last  time  in  the  Siwaliks  of  Perim  Island,  Kach,  Sind,  and  the 
Punjab ;  while  the  mastodons  include  a  large  number  of  species, 
some  of  which  present  such  a  close  approximation  to  the  so-called 
stegodont  elephants  (which,  as  already  mentioned1,  are  peculiar  to 
this  fauna)  as  to  render  it  impossible  to  draw  any  well-defined 
demarcation  between  the  genera  Mastodon  and  Elephas.  Not 
only  does  the  Siwalak  fauna  include  the  aforesaid  stegodont,  or 

1  Stipra,  p.  172. 


V.]  S1WALIK   FAUNA.  2O; 

transitional  elephants,  but  likewise  one  which  may  well  have  been 
the  ancestor  of  the  species  now  inhabiting  India.  Eastwards  these 
transitional  elephants  and  mastodons  have  been  traced  into  Java, 
Borneo,  China,  and  Japan  ;  and,  as  stated  in  an  earlier  chapter, 
there  can  be  no  doubt  that  the  modern  elephants  were  evolved  in 
this  area. 

Although,  as  shown  in  the  foregoing  survey,  the  Siwalik  fauna 
differs  in  certain  respects  from  that  of  Pikermi,  Samos,  Leberon, 
etc.,  yet  there  can  be  no  hesitation  in  regarding  the  whole  lower 
Pliocene  fauna  of  Europe,  North  Africa,  Asia  Minor,  and  South 
and  East  Asia  as  essentially  one ;  and  consequently  at  this  epoch 
there  was  no  possibility  of  distinguishing  between  the  Palaearctic 
and  Oriental  regions.  Whence  Ethiopian  Africa  had  by  this 
time  received  the  forerunners  of  its  present  higher  mammalian 
fauna,  we  have,  unfortunately,  no  decisive  evidence.  Writing 
some  years  ago,  Dr  Blanford1  seems  to  suggest  that  the  irruption 
of  the  modem  African  fauna  was  anterior  to  the  Pliocene.  After 
referring  to  certain  peculiarities  connected  with  the  existing 
mammalian  fauna  of  India  and  the  Malayan  area,  he  observes  that 
"  these  cases  of  isolation  probably  indicate  that  the  animals  belong 
to  an  older  fauna,  now  partly  replaced  by  newer  types,  and  that 
the  older  fauna  was  common  to  India  and  Africa.  It  is  very 
probable  that  these  animals  are  descended  from  the  ancient 
tropical  fauna  of  the  early  Tertiary  times.  But,  so  far  as  it  is 
possible  to  judge,  the  process  of  variation  would  have  caused  a 
greater  distinction  between  forms  so  widely  separated  and  exposed 
to  such  different  conditions,  if  the  period  of  isolation  were  great ; 
and  it  is  difficult  to  suppose  that  the  lands  inhabited  by  the 
ancestors  of  the  Simiidce,  Lemuridce,  Tragulida,  and  Manidce  of 
the  Oriental  and  Ethiopian  regions  can  have  been  separated  prior 
to  the  early  part  of  the  Miocene  period." 

This  is  perfectly  true  so  far  as  it  goes,  but  since,  as  we  have 
seen,  genera  like  Hippopotamus,  Bos,  Capra,  Equus,  and  Elephas 
are  unknown  previous  to  the  Siwalik  epoch,  and  some  of  them 
at  least  were  evolved  at  or  about  that  time  in  the  Indian  area,  it 
seems  necessary  to  assume  the  existence  of  a  free  land  communi- 

1  Manual  of  Geology  of  India,  ist  ed.  p.  Ixviii.  (1879). 


208  EASTERN   ARCTOG^A.  [CHAP. 

cation  between  the  Ethiopian  and  Oriental  regions  at  least  as  late 
as  the  lower  Pliocene  epoch.  With  regard  to  where  this  connec- 
tion was  situated,  we  may  note,  in  the  first  place,  that  Dr  Wallace : 
was  of  opinion  that  even  the  Pikermi  fauna  made  its  way  into 
Africa  chiefly  through  Syria,  although  a  brief  connection  of  Europe 
with  Tunis  is  admitted.  When  the  passage  in  question  was 
written,  little  or  nothing  was,  however,  known  as  to  the  Pliocene 
fauna  of  Algeria.  And  although  this  undoubtedly  indicates  a 
western  connection  between  Europe  and  Africa,  yet  even  in  the 
Pliocene  the  Sahara  probably  formed,  as  now,  a  barrier2  across 
which  the  fauna  of  northern  Africa  could  not  pass  south.  Accord- 
ingly, even  the  Pikermi  fauna  may  have  come  round  byway  of  Egypt. 
Be  this  as  it  may,  it  seems  clear  that  the  Siwalik  fauna  entered 
Africa  by  way  of  Syria  or  Arabia,  or  possibly  by  both.  The  most 
direct  line  of  communication  would  be  via  the  Gulfs  of  Oman  and 
Aden ;  and  some  indication  that  such  a  line  of  connection  may 
have  existed  is  afforded  by  the  distribution  of  the  goats  of  the 
genus  Hemitragus.  As  already  stated,  fossil  species  of  this  genus 
occur  in  the  Siwaliks  of  Perim  Island  and  the  Himalaya,  while  of 
the  three  existing  forms,  one  is  Himalayan,  a  second  confined  to 
the  Nilgiri  and  certain  other  South  Indian  ranges,  and  the  third 
inhabits  Oman.  So  far  as  it  goes,  the  evidence  of  these  goats  is 
strongly  suggestive  of  the  former  existence  of  a  land-bridge  across 
the  mouth  of  the  Persian  Gulf,  as  otherwise  we  should  expect  to 
find  living  species  in  Persia  and  other  parts  of  western  Asia.  If 
the  existence  of  such  a  bridge  be  admitted,  we  only  require  another 
across  the  narrow  strait  of  Bab-el-Mandeb  to  give  a  free  line  of 
communication  between  India  and  Africa  in  this  direction. 

Whether,  however,  the  migration  from  India  to  Africa  took 
place  at  the  north  or  south  end  of  the  Red  Sea,  or  at  both  ends, 
it  is  certain  that  the  connecting  land  must  have  been  of  consider- 
able width,  and  suited  to  the  passage  of  mammals  of  all  kinds. 
In  referring  to  the  nature  of  the  connection,  Dr  Wallace3  remarks 
that  "  we  may  now  perhaps  see  the  reason  of  the  singular  absence 

1  Geographical  D^strib^ttwn  of  Animals,  Vol.  I.  p.  288. 

2  The  idea  that  there  was  a  Tertiary  sea  in  the  Sahara  is  incorrect;  see 
Blanford,  Quart,  Journ.  GeoL  Soc.  Vol.  XLVI.  p.  90  (1890). 

3  Op.  cii.  p.  291. 


V.]  SIWALIK   FAUNA.  209 

from  tropical  Africa  of  deer  and  bears ;  for  these  are  both  groups 
which  live  in  fertile  or  well-wooded  countries,  whereas  the  line  of 
immigration  from  Europe  to  Africa  was  probably  always,  as  now, 
to  a  great  extent  a  dry  and  desert  tract,  suited  to  antelopes  and 
large  felines,  but  almost  impassable  to  deer  and  bears."  The 
Siwalik  chimpanzee,  however,  indicates  most  unmistakably  that 
the  communication  by  way  of  Arabia  or  Syria  between  the  Ethio- 
pian and  Oriental  regions  must  have  embraced  a  forest-area,  and 
accordingly  have  been  of  considerable  width. 

With  regard  to  the  question  why  so  many  genera  which  existed 
in  India  and  southern  Europe  during  the  Pliocene  should  have 
disappeared  from  those  areas  to  live  on  in  Africa,  all  we  can  say  is 
that  it  is  quite  evident  that  a  southern  migration  of  the  fauna  has 
certainly  taken  place,  and  that  this  was  probably  induced  by  the 
cold  heralding  the  approach  of  the  glacial  period.  Although  we 
have  few,  if  any,  decisive  physical  evidences  of  a  cold  period  in 
India,  yet  the  existence  of  a  goat  (Hemitragus)  nearly  allied  to  a 
Himalayan  species  in  the  ranges  of  southern  India  seems  to  indi- 
cate that  such  must  have  occurred,  as  it  would  be  quite  impossible 
for  the  ancestral  form  to  have  crossed  the  intervening  plains  under 
present  conditions  of  temperature.  It  is  further  noteworthy  that 
many  of  the  animals  which  have  disappeared  from  India,  such 
as  chimpanzees,  hippopotami,  giraifes,  water-chevrotains,  and 
ostriches,  are  precisely  those  which  are  now  restricted  to  very  hot 
climates  ;  whereas  the  lion,  tiger,  rhinoceroses,  elephants,  and 
monkeys,  which  both  now  or  during  the  Plistocene  are  known 
to  be  capable  of  existing  in  cold  climates,  have  persisted. 

Leaving  these  exceedingly  difficult  questions,  two  other  points 
may  be  noticed  in  connection  with  the  Siwalik  fauna.  In  the 
first  place,  since  the  Siwalik  hills  themselves  form  ranges  of  con- 
siderable height  on  its  southern  flank,  it  is  evident  that  the 
Himalaya  was  much  lower  during  the  lower  Pliocene  epoch  than 
it  is  at  present ;  Dr  Blanford l  stating  that  the  movement  which 
led  to  its  elevation  "has  been  distributed  over  the  Tertiary  and 
post-Tertiary  period,  and  a  great  portion  in  post-Plistocene." 
This  will  account  for  the  community  between  the  lower  Pliocene 

1  See  Geol.  Mag.  Decade  3,  Vol.  ix.  p.  166,  note  (1892). 
L.  14 


210  EASTERN   ARCTOG^EA.  [CHAP. 

fauna  of  the  Himalayan  area  and  Mongolia,  the  Himalaya  at  that 
epoch  not  forming,  as  now,  an  impassable  barrier  to  the  north  of 
the  Oriental  region.  The  second  point  relates  to  the  survival  in 
the  Siwalik  fauna  of  archaic  forms,  which  had  disappeared  at  that 
date  from  Europe.  This  fact,  especially  since  old  types  such  as 
lemurs  and  gymnuras  are  even  now  met  with  in  the  Oriental 
region,  lends  support  to  the  view  advanced  in  an  earlier  chapter1 
that  marsupials  may  have  lived  on  in  south-eastern  Asia  long  after 
they  had  completely  disappeared  from  Europe. 

Our   knowledge    of    the    later    Pliocene   faunas    of    Eastern 

Arctogaea  is  mainly  confined  to  Europe,  where  at 
cene Kaunas.0"  this  period  the  general  distribution  of  land  and  sea 

was  apparently  very  much  the  same  as  at  the 
present  day.  Spain  was,  however,  connected  with  Africa,  as  was 
probably  also  Italy  by  way  of  Sardinia  and  Malta.  A  portion  of 
Italy  was,  however,  submerged,  while  in  Belgium,  Holland,  and 
the  south-east  of  England  the  sea  intruded  upon  what  is  now  land ; 
but,  on  the  other  hand,  Britain  was  joined  to  the  Continent. 
Few  mammaliferous  deposits  of  this  age  have  been  preserved  to 
us,  but  among  these  are  the  Crags  of  the  east  coast  of  England 
(which  contain  numerous  fossils  derived  from  earlier  formations), 
the  fresh-water  beds  of  the  Val  d'  Arno  in  Italy,  as  well  as  others  in 
the  Auvergne,  in  the  Rhone  valley,  at  Roussillon,  and  in  the 
neighbourhood  of  Montpellier.  The  following  genera  are  in- 
cluded in  this  fauna,  those  which  are  extinct  having  an  asterisk 
prefixed. 

PRIMATES.          Semnopithecus. 

*  Dolichopithecus. 
Macacus. 

INSECTIVORA.     Sorex  (Shrews). 
CARNIVORA.     *Machaerodus  (Sabre-tooths). 

Felis  (Cats). 

Viverra  (Civets). 

Hyaena. 

Canis  (Wolves  and  Foxes). 

*  Hyaenarctus. 

1  Supra,  p.  57. 


v.] 


LATER   PLIOCENE   FAUNAS. 


211 


CARNIVORA  (cont.). 

Ursus  (Bears). 

^Elurus  (Cat-bears). 

Mustek  (Martens  and  Weasels). 

Lutra  (Otters). 
RODENTIA.          Arctomys  (Marmots). 

*  Chalicomys. 
Castor  (Beaver). 

*Trogontherium  (Giant  Beaver). 

Cricetus  (Hamsters). 

Microtus  (Voles). 

Mus  (Rats  and  Mice). 

Hystrix  (Porcupines). 
*Pellegrinia, 
*Myolagusj 

Lagomys  J 

Lepus  (Hares) 

Sus  (Pigs). 

Hippopotamus. 

Cervus  (Deer). 

Alces  (Elk). 

Cervulus  (Muntjacs). 

*  Palaeoryx. 
Gazella  (Gazelles). 
Bos  (Oxen). 
Tapirus  (Tapirs). 
Rhinoceros. 
Equus  (Horses). 

*Hipparion — very  rare. 

*  Mastodon! 


UNGULATA. 


Elephas    J 


(Elephants). 


In  this  list  by  far  the  greater  number  of  the  genera  are  living 
ones,  and  if  we  removed  from  it  types  like  Hyana,  Hippopotamus, 
Rhinoceros  and  Elephas,  which  were  spread  during  the  Pliocene 
and  Plistocene  epochs  over  the  greater  part  of  Eastern  Arctogaea, 
its  Ethiopian  resemblances  are  by  no  means  strongly  marked. 
Although  the  larger  forms  (as  in  the  succeeding  Plistocene  epoch) 

14—2 


212  EASTERN    ARCTOG^A.  [CHAP.  V. 

include  a  considerable  number  of  genera  now  mainly  confined  to 
tropical  or  subtropical  countries,  the  rodent  fauna  exhibits  a 
marked  Palsearctic  facies,  thus  indicating  an  approximation  to  the 
existing  state  of  things.  Among  the  extinct  rodents,  Pellegrinia, 
from  the  Sardinian  Pliocene,  belongs,  however,  to  the  Octodontida, 
and  is  probably  allied  to  the  existing  African  Ctenodactylus.  Tro- 
gontherium  is  a  gigantic  extinct  type  of  beaver,  which  also  persisted 
into  the  Plistocene.  The  deer  include  northern  types  unknown 
in  the  lower  Pliocene. 

One  of  the  most  remarkable  features  of  this  fauna  is  the 
occurrence  of  a  large  species  of  sElurus, — a  genus  represented 
elsewhere  only  by  the  cat-bear  or  panda  (^E.  fulgens)  of  the 
eastern  Himalaya,  which,  although  formerly  regarded  as  the  type 
of  a  family  by  itself,  is  now  included  in  the  American  Procyonidce 
(raccoons).  The  fossil  species  has  been  hitherto  detected  only  in 
the  English  Crag ;  the  genus  may,  however,  be  expected  to  occur 
in  the  Siwaliks,  since  it  is  quite  clear  that  it  must  have  been 
originally  connected  with  the  American  representatives  of  the 
family  by  forms  inhabiting  Eastern  Asia. 

With  the  end  of  the  Pliocene  epoch  this  brief  survey  of  the 
Tertiary  mammalian  faunas  of  Eastern  Arctogaea  may  be  brought 
to  a  close,  since  the  Plistocene  mammals  can  be  more  con- 
veniently considered  under  the  headings  of  the  different  regions  of 
this  great  province.  While  throughout  the  Oligocene,  Miocene, 
and  lower  Pliocene  epochs  no  trace  of  the  present  zoological 
regions  of  this  half  of  the  Arctogseic  realm  is  shown,  when  the 
Upper  Pliocene  is  reached  there  are  faint  indications  of  the 
demarcation  of  the  Eastern  Holarctic.  At  the  time  of  the  Plisto- 
cene, as  will  be  shown  in  a  later  chapter,  the  Eastern  Holarctic, 
Oriental  and  Ethiopian  regions  appear  to  have  assumed  a  still 
more  marked  distinction,  although  this  is  to  a  great  extent 
obscured  by  the  wide  range  even  at  that  epoch  of  genera  like 
Hippopotamus,  Rhinoceros,  Elephas,  Macacus,  etc.  Moreover, 
several  species  which  are  now  confined  to  one  of  the  three  regions 
in  question  had  then  a  more  extensive  distribution,  so  that  it  is 
only  during  the  recent  epoch  that  the  Holarctic,  Oriental,  and 
Ethiopian  regions  attained  the  full  faunistic  peculiarities  by  which 
they  are  now  characterised. 


CHAPTER  VI. 

THE   MALAGASY   REGION. 

Limits — Mammalian  Fauna — Relations  of  Madagascar  to  the  Mainland. 

INCLUDED  by  Drs  Sclater  and  Wallace  within  the  Ethiopian 
region,  Madagascar  and  the  adjacent  groups  of 
islands  were  referred  to  a  region  apart  by  Dr  Blan- 
ford1;  this  separation  being  justified  not  only  by  the  mammalian 
fauna,  but  likewise  by  many  other  groups  of  animals.  To  quote 
Dr  Wallace,  this  region  "comprises,  besides  Madagascar,  the 
islands  of  Mauritius,  Bourbon,  and  Rodriguez,  the  Seychelles, 
and  Comoro  Islands.  Madagascar  itself  is  an  island  of  the 
first  class,  being  a  thousand  miles  long,  and  about  two 
hundred  and  fifty  miles  in  average  width.  It  lies  parallel  to 
the  coast  of  Africa,  near  the  southern  tropic,  and  is  separated 
by  230  miles  of  sea  from  the  nearest  part  of  the  continent, 
although  a  bank  of  soundings  projecting  from  its  western 
coast  reduces  this  distance  to  about  160  miles.  Madagascar  is  a 
mountainous  island,  and  the  greater  part  of  the  interior  consists  of 
open  elevated  plateaus ;  but  between  these  and  the  coast  there 
intervene  broad  belts  of  luxuriant  tropical  forests."  It  is  this 
forest-district  which  forms  the  home  of  most  of  its  peculiar  fauna. 
As  regards  geological  structure,  it  [appears  from  the  researches  of 
Messrs  Cortese  and  Baron  that,  roughly  speaking,  a  line  drawn 
from  north  to  south  so  as  to  divide  the  island  into  two  longitudinal 
halves,  gives  an  area  of  granitic  and  volcanic  rocks  on  the  right  or 
eastern  side,  and  on  the  left  or  western  side  one  of  sedimentary 
deposits,  containing  beds  belonging  to  the  Jurassic,  Cretaceous, 
Eocene  and  recent  epochs.  Blown  sand  occurs  in  abundance 

1  Appendix,  No.  8,  p.  76. 


214  THE   MALAGASY   REGION.  [CHAP. 

around  the  coast,  and  numerous  old  lake-basins  or  marshes,  some 
of  very  large  dimensions,  form  receptacles  where  remains  of  the 
later  faunas  have  been  preserved.  With  the  exception  of  the 
Comoro  group,  which  contain  a  few  species,  the  non-volant  mam- 
malian fauna  is  confined  to  Madagascar,  so  that  the  other  islands 
do  not  properly  come  within  the  province  of  the  present  work. 
It  is,  however,  important  to  observe  that  the  Seychelles  differ  from 
almost  all  oceanic  islands  in  consisting  largely  of  granitic  and 
other  crystalline  rocks. 

In  an  island  lying  so  close  to  the  African  continent  as  Mada- 
gascar, the  natural  assumption  would  be  that,  if  it 
fa^ia"1"  possessed  a  mammalian   fauna  at  all,   such  fauna 

would  be  closely  allied  to  that  of  the  mainland.  As 
a  matter  of  fact,  precisely  the  reverse  is  the  case,  and  out  of  a 
total  of  fully  28  genera  of  non-volant  mammals  now  or  recently 
inhabiting  the  island,  only  three  are  common  to  Africa.  This, 
however,  is  by  no  means  all,  for  out  of  these  three  genera  two 
{Hippopotamus  and  Sus]  are  such  as  have  probably  crossed  the 
intervening  channel,  although  at  a  time  when  it  was  narrower  than 
at  present,  while  it  is  quite  possible  that  the  third  (Crocidura)  may 
have  been  introduced  by  human  agency.  Even  this,  however, 
scarcely  gives  a  true  idea  of  the  case.  In  the  first  place,  not  only 
are  the  peculiar  genera  unknown  in  Africa,  but  they  are  equally 
strange  to  all  the  other  regions  of  the  world.  In  the  second  place, 
these  genera  belong  to  groups  which  form  only  a  very  small  por- 
tion of  the  existing  mammalian  fauna  of  the  Ethiopian  region. 
At  the  present  day,  as  will  be  more  fully  indicated  in  the  following 
chapter,  Ethiopian  Africa  is  especially  characterised  by  its  nume- 
rous antelopes,  as  well  by  giraffes,  zebras,  rhinoceroses,  elephants, 
hippopotami,  wart-hogs,  bush-pigs,  lions,  leopards  and  various 
other  large  cats,  baboons,  anthropoid  apes,  aard-varks,  and 
ostriches.  But,  with  the  exception  of  the  aforesaid  bush-pig  and 
extinct  hippopotamus,  not  a  single  representative  of  any  one  of 
these  groups  is  found  in  Madagascar.  In  place  of  such  animals, 
Madagascar  is  populated  by  a  host  of  lemurs,  so  numerous  that 
the  number  of  their  species  considerably  exceeds  that  of  all  the 
other  non-volant  mammalian  inhabitants  of  the  island.  Civet- 
and  mungoose-like  species,  all  pertaining  to  peculiar  genera,  alone 


VI.]  THE   FAUNA.  21$ 

represent  the  numerous  Garni vora  of  the  mainland;  the  Insectivora, 
in  addition  to  the  aforesaid  Crocidura,  or  musk-shrew,  include  only 
the  peculiar  family  of  the  tenrecs  (Centettdce),  which  is  confined  to  the 
island,  and  a  representative  of  the  Ethiopian  family  Potamogalida ; 
while  the  rodents  comprise  five  genera  of  the  cosmopolitan 
mouse-family  (Muridce],  more  or  less  closely  allied  to  one  another, 
but  different  from  any  found  elsewhere. 

The  following  is  a  list  of  the  genera  of  non-volant  Malagasy 
mammals ;  those  which  are  extinct  being  indicated  by  an  asterisk, 
and  the  names  of  all  the  groups  peculiar  to  the  island  printed  in 
italics. 

PRIMATES. — LEMUROIDEA. 
LEMURID^E. 

Chirogale  (Mouse-lemurs) ;  4  species.  £jieiroAd.le<*s 

Microcebus  (Dwarf  Lemurs) ;  5  species.    *• 

Opolemur  (Fat-tailed  Lemurs) ;  2  species.  ^ 

Lemur  (True  Lemurs) ;  8  species. 

Mixocebus  (Hattock) ;   i  species.  ?^ 

Hapalemur  (Gentle  Lemurs) ;  2  species.  'y  (     \ 

Lepidokmur  (Sportive  Lemurs) ;  8  species.     7.'  $c»~*V\ 

Avahis  (Avahi) ;   i  species.  ? 

Propithecus  (Sifakas) ;  4  species.^7- 

/^m'(Endrina) ;   i  species. 

*  ME  GAL  AD  A  PID^E. 

*  Megaladapis  (Giant  Lemur);  i  species.         K««^l»*< 
CHIROMYID&.      t\ 

$J±fL 

Chiromys  (Aye-aye) ;   i  species. 
INSECTIVORA. 
SORICID.E. 

Crocidura  (Musk-shrews) ;  i  species. 
CENTETID^E. 

Centetes  (Tenrec) ;  i  species. 

Hemicentetes  \  2  species. 

Ericulus  (Hedgehog-tenrec) ;  i  species. 

Echinops-j  i  species. 

Microgale  (Long- tailed  tenrecs) ;  3  species. 

Oryzorictes  (Rice-tenrecs) :  2  species. 


2l6  THE   MALAGASY  REGION.  [CHAP. 

INSECTIVORA  (cent.). 

POTAMOGALID^E. 

Geogale;  i  species. 
CARNIVORA. 

VlVERRID^E. 

CryptoproctincR. 

Cryptoprocta  -,  i  species. 

Viverrinae. 

Fossa  ;  i  species. 
Herpestinse. 

Galidictis  (Striped  Mungooses)  ;  2  species. 

Galidia  (Ring-tailed  Mungoose)  ;  i  species. 

Hemigalidia  (Brown-tailed  Mungoose)  ;  i  species. 
Euplerince. 

Eupleres  (Small-toothed  Mungoose)  ;  i  species. 

RODENTIA. 

MURID/E. 

Hypogeomys  ;   i  species. 
Nesomys  ;  2  species. 
Brachytarsomys  ;   i  species. 
Hallomys;  i  species. 
Eliurus;  2  species. 

UNGULATA. 


Sus  (Potamochcerus)  ;   t  species. 

HIPPOPOTAMID^:. 

Hippopotamus  ;  i  species  (extinct). 

Considering  the  fauna  in  more  detail,  it  may  be  first  mentioned 
that  the  lemurs  (Lemuroidea)  differ  from  the  higher,  or  Anthropoid 
Primates  by  their  generally  lower  grade  of  organisation,  as  well  as 
by  certain  features  of  the  skull  and  internal  anatomy  which  need 
not  be  more  fully  noticed  here.  They  all  have  fox-like,  expression- 
less faces  ;  and,  with  the  exception  of  the  aye-aye  and  the  Asiatic 
tarsiers,  they  are  characterised  by  the  innermost  pair  of  upper 
incisor  teeth  being  separated  from  one  another  in  the  middle  line. 


VI.]  LEMUROIDS.  217 

At  the  present  day  lemuroids  are  represented  elsewhere  only 
in  the  Ethiopian  and  Oriental  regions ;  the  African  forms  being 
more  nearly  allied  to  the  Malagasy  types  than  are  those  of  Asia. 
As  stated  in  an  earlier  chapter,  the  group  was,  however,  well 
represented  in  the  lower  Oligocene  of  western  Europe,  where 
certain  forms  (Mtcroc/ioerus)  distinctly  approximate  some  of  the 
living  kinds,  although  differing  in  the  conformation  of  the  first 
lower  premolar  tooth,  which  in  the  existing  Lemuridce,  assumes 


FlG.  48.      SKULL   OF    LEMUR. 

tic.  upper  canine  ;  k.  lower  canine ;  pm.  premolars ;  m.  molars. 

the  form  and  function  of  a  canine  or  tusk.  In  the  latter  family  (of 
which  the  distribution  is  coextensive  with  that  of  the  suborder)  the 
first  three  genera  in  the  foregoing  list  belong  to  a  subfamily 
(Galagina)  distinguished  by  the  elongation  of  the  bones  of  the 
tarsus,  and  represented  by  an  allied  genus  (Galago)  on  the  African 
mainland.  The  next  four  genera  constitute  the  typical  subfamily 
(Lemurincz\  which  is  absolutely  confined  to  Madagascar  and  some 
of  the  islands  of  the  Comoro  group,  and  of  which  the  ring-tailed 
lemur  (Lemur  cattd)  is  one  of  the  most  familiar  examples  in 
European  menageries.  All  these  lemurs,  which  have  long, 
although  non- prehensile  tails,  differ  from  the  first  subfamily  by  the 
normal  structure  of  the  bones  of  the  ankle.  The  third  subfamily 
(Indrisin<z\  which  is  likewise  peculiar  to  this  region,  includes  the 
avahi,  sifakas,  and  the  endrina,  all  of  which  differ  from  the  two 
preceding  groups  by  having  only  thirty,  in  place  of  thirty-six  teeth ; 
while  the  endrina  is  peculiar  in  having  the  tail  rudimentary.  The 
group  includes  the  largest  living  lemurs ;  the  sifakas  and  endrina 


218 


THE    MALAGASY   REGION. 


[CHAP. 


differing  from  other  members  of  the  suborder  by  their  diurnal 
habits.  They  form  a  characteristic  feature  in  every  wooded  Mala- 
gasy landscape,  there  being  scarcely  a  copse  in  the  island  which  is 
not  tenanted  by  one  or  more  of  these  strange  creatures ;  and  when 


FIG.  49.     RING-TAILED  LEMUR  (Lemur  catta). 

walking  from  covert  to  covert,  they  do  so  in  an  erect  posture,  with 
their  hands  clasped  behind  their  necks. 

Whereas  the  endrina  (the  largest  living  lemur)  is  only  two  feet 
in  length  exclusive  of  the  rudimentary  tail,  the  extinct  Megala- 
dapis,  whose  remains  have  been  obtained  from  the  Ambolisatra 
marsh,  had  a  skull  three  times  the  size  of  that  of  the  latter,  so  that 


VI.]  LEMUROIDS.  2IQ 

the  whole  animal  might  be  compared  in  size  to  a  mandrill.  The 
skull  of  this  species  is  characterised  by  the  great  elongation  of 
the  face,  and  in  several  respects  shows  resemblances  to  that 
of  the  European  Oligocene  genus  Adapts]  although  the  upper 
molar  teeth  are  peculiar  in  having  tritubercular  crowns,  whereas 
those  of  all  modern  lemurs  are  quadrangular.  There  is  consider- 
able reason  to  believe  that  the  giant  lemur  was  actually  living  in 
the  middle  of  the  seventeenth  century,  an  otherwise  unknown 
animal  being  described  by  De  Flacourt  in  1658  under  the  name  of 
tretretretre,  or  tratratratra,  which  accords  fairly  well  with  the  fossil 
remains.  The  giant  lemur  is,  however,  not  the  sole  extinct 
member  of  the  group  from  Madagascar,  since  the  hinder  part  of  a 


FlG.    50.      SKULL    OF   AYE- AYE. 

skull  indicates  another,  but  at  present  unnamed  genus,  apparently 
allied  to  Hapalemur.  The  last  of  the  Malagasy  lemurs  is  the 
singular  aye-aye  (Chiromys);  a  creature  representing  by  itself  a 
separate  family,  broadly  distinguished  from  all  other  members  of 
the  suborder  by  the  curious  resemblance  of  the  dentition  to  that  of 
the  rodents,  to  say  nothing  of  the  extreme  elongation  and  slender- 
ness  of  the  middle  finger  of  the  hand. 

Apart  from  the  single  musk-shrew,  the  Malagasy  insectivores 
all  belong  to  the  group  with  tritubercular  upper  molar  teeth, 
which,  as  already  mentioned,  is  now  confined  to  the  more  southern 
portions  of  the  world,  and  is  evidently  a  very  primitive  one.  The 
small  mouse-like  creature  (Geogale  auritd]  representing  the  Pota- 


220  THE   MALAGASY   REGION.  [CHAP. 

mogalidce  differs  from  its  Ethiopian  cousin,  not  only  in  its  inferior 
dimensions,  but  likewise  in  having  but  thirty-four  in  place  of  forty 
teeth ;  and  it  is  possible  that,  when  more  fully  known,  it  will  have 
to  be  assigned  to  a  family  by  itself.  As  stated  in  a  previous 
chapter1,  the  tenrecs  (Centetida)  appear  to  have  their  nearest  allies 
in  the  West  Indian  solenodons,  although  the  relationship  is  now 
believed  to  be  somewhat  less  close  than  was  formerly  supposed. 
The  common  tenrec  ( Centetes),  which  is  the  largest  member  of  its 
order,  measuring  from  a  foot  to  sixteen  inches  in  length,  is  a  tail- 
less creature,  remarkable  for  the  possession  of  four  pairs  of  upper 
molar  teeth,  as  in  marsupials.  Much  smaller  are  the  two  species 
of  Hemicentetes ,  which,  in  addition  to  differences  in  the  dentition, 
are  distinguished  by  having  rows  of  spines  along  the  back  at  all 
ages,  instead  of  merely  in  the  young  condition.  The  hedgehog- 
tenrecs,  forming  the  genera  Ericulus  and  Echinops,  are  small 
forms  having  the  whole  of  the  back  and  tail  covered  with  close-set 
spines.  The  two  other  genera  are  spineless  at  all  ages ;  Microgale 
being  readily  distinguished  by  the  inordinate  length  of  the  tail 
which  is  equal  to  twice  that  of  the  head  and  body,  while 
Oryzorictes  has  this  appendage  relatively  short. 

The  largest,  and  at  the  same  time  one  of  the  most  peculiar  of 
the  Malagasy  carnivores  is  the  fossa  ( Cryptoprocta),  which,  although 
usually  included  in  the  Viverridce,  is  so  different  from  all  other 
members  of  that  group  that  it  has  been  regarded  as  constituting  a 
family  by  itself,  specially  characterised  by  the  feline  type  of  denti- 
tion. On  the  other  hand,  Daubenton's  civet  (Fossa),  although 
representing  a  genus  by  itself,  has  its  nearest  relative  in  the  widely 
distributed  Oriental  rasse  ( Viverra  malaccensts).  The  latter  species, 
although  now  found  both  in  Madagascar  and  the  Comoro  group, 
has  in  all  probability  been  introduced  there.  Of  the  four  remain- 
ing genera,  Galidictis,  Galidia,  and  Hemigalidia  are  more  or  less 
closely  allied  to  the  mungooses,  although  presenting  certain 
structural  differences  from  other  genera ;  but  the  fourth  (Eupleres) 
is  so  markedly  distinct  as  to  constitute  a  subfamily  by  itself. 

The  five  genera  of  murine  rodents  call  for  but  little  remark, 
although  it  is  noteworthy  that  they  are  all  more  or  less  closely 

1  Supra,  p.  70. 


VI.] 


CARNIVORA. 


221 


allied,  and  belong  to  the  cricetine  section,  which  contains  the 
oldest  members  of  the  family.  Nothing  need  be  said  in  regard  to 
the  two  ungulates,  except  that  they  both  belong  to  Ethiopian 
types.  Although  bats  are  not  taken  much  into  account  in  the 
present  volume,  it  is  important  to  notice  a  peculiar  distribution  of 
the  fruit-bats  or  Pteropodida ;  more  especially  as  this  coincides  with 
that  of  many  Malagasy  birds.  On  this  point  Dr  Blanford  writes 
that  "  the  only  African  genus  belonging  to  the  family  is  Epomo- 


FIG.  51.     THE  FOSSA  ( Cryptoprocta  ferox] . 

^^  which  is  confined  to  the  continent,  whilst  throughout  the 
Mascarene  archipelago,  and  even  in  the  Comoro  islands  in  the 
Mozambique  channel,  the  typically  Oriental  genus  Pteropus  occurs, 
and  is  represented  in  various  islands  by  five  species,  one  or  two  of 
them  only  distinguished  by  critical  characters  from  the  common 
'  flying-fox '  of  the  Indian  peninsula." 

In    groups    other   than    mammals,    certain    common   features 
between  the  reptiles  of  Madagascar  and  South  America  have  been 

1  A  second  genus,  Scotonycteris,  has  been  described  from  the  Cameruns 
since  this  passage  was  written  ;  and  the  author  has  omitted  mention  of 
Trygenycferis  (Megaloglossus}. 


222  THE    MALAGASY   REGION.  [CHAP. 

mentioned  in  an  earlier  chapter1,  where  it  was  attempted  to  show 
that  although  these  instances  of  discontinuous  distribution  might 
be  explained  by  parallel  migration  from  a  common  northern 
centre,  yet  that  the  Tertiary  mammalian  evidence  indicated  that 
the  American  forms  had  reached  their  present  habitat  by  way  of 
Madagascar  and  Africa.  It  will  suffice  to  add  here  that  giant 
land-tortoises,  which  existed  in  the  Mascarenes  during  the  present 
epoch,  are  represented  by  extinct  species  from  the  superficial 
deposits  of  Madagascar;  and  that  the  latter  have  also  yielded 
remains  of  gigantic  flightless  birds  (dEpyornis)  markedly  distinct 
from  any  other  known  type.  And  here  it  may  be  mentioned  that 
the  chamseleons  (Chamaleontida)  present  a  certain  similarity  in 
their  distribution  to  the  lemuroids,  the  Malagasy  region  including 
23  out  of  the  49  species,  while  nearly  all  the  others  are  Ethiopian. 
As  a  whole,  the  Malagasy  reptiles,  with  the  exception  of  the 
snakes,  are  stated  to  be  more  nearly  allied  to  those  of  the  main- 
land than  are  either  the  mammals  or  the  birds ;  but  the 
amphibians  exhibit  more  decided  traces  of  Oriental  affinities. 

Concentrating  our  attention  mainly  on  the  mammals  alone, 
their  distinctness  from  those  of  all  other  parts  of 

Relations  of  .... 

Madagascar  to     the  world  are  quite  sufficient  to  indicate  the  right  of 

the  Mainland.        _  _     ,  r  -\  r  ~\       • 

Madagascar  to  form  the  centre  of  a  separate  zoologi- 
cal region.  In  the  survey  of  the  lower  Oligocene  fauna  of  Europe 
it  has  been  shown  that  both  lemuroids  and  civet-like  carnivores 
were  common,  one  of  the  latter  having  been  referred  to  the  existing 
genus  Viverra.  Hence  it  is  probable  that  to  this  fauna  we  must 
look  for  the  ancestors  of  the  Malagasy  mammals.  The  only 
lemuroids  closely  allied  to  those  of  Madagascar  are  the  African 
galagos,  and  as  the  civet-family  ( Viverridcz]  is  better  represented 
in  Africa  than  elsewhere,  it  may  be  taken  for  granted  that  Mada- 
gascar received  its  mammalian  fauna  from  the  mainland.  Putting 
aside  the  hippopotamus  and  bush-pig,  which  doubtless  arrived 
later,  the  Malagasy  fauna  can,  however,  have  been  derived  from 
Africa  only  at  a  time  anterior  to  the  introduction  of  the  modern 
types  of  ungulates  into  that  continent,  when  it  was  chiefly  popu- 
lated by  lemuroids  and  civet-like  carnivores2.  The  question  then 

1  Supra,  p.  131. 

'2  It  is  of  course  probable  that  some  of  the  Oligocene  primitive  ungulates 


VI.]  RELATION    TO   AFRICA.  223 

narrows  itself  as  to  the  probable  date  of  the  connection  between 
the  island  and  the  continent.  Now,  so  far  as  can  be  determined, 
none  of  the  European  Oligocene  lemuroids  are  referable  to  the 
family  Lemuridce  •  and  since  both  the  Ethiopian  and  Malagasy 
representatives  of  the  subfamily  Galagince  resemble  one  another  in 
the  peculiar  structure  of  the  ankle,  or  tarsus,  it  is  pretty  evident 
that  not  only  was  the  family,  but  likewise  the  subfamily  differenti- 
ated before  the  separation  of  Madagascar.  Allowing  time  for  the 
southward  migration  of  the  Oligocene  lemuroids  and  civets,  arid 
the  modification  of  the  former  into  the  Galagince,  it  seems 
impossible  to  put  the  separation  at  an  earlier  date  than  the  Upper 
Oligocene,  while  it  might  well  be  Miocene1.  Confirmation  of 
this  comparatively  late  separation  of  the  island  is  afforded  by  some 
observations  of  Dr  Blanford  with  regard  to  the  passage  of  the 
bush-pig  across  the  intervening  strait,  for  it  is  evident  that  both 
that  animal  and  the  hippopotamus  must  have  reached  Madagascar 
by  swimming,  as  otherwise  more  ungulates  would  assuredly  have 

migrated  into  Africa  with  the  lemuroids  and  Viverridce ;  but  if  so,  all  have 
died  out.  The  Tertiary  palseontological  history  of  Africa  or  Madagascar  can 
alone  decide  this  point ;  but  if  ancestors  of  the  South  American  extinct  ungu- 
lates reached  their  home  by  way  of  Africa,  it  is  certain  that  primitive  members 
of  that  order  must  have  first  passed  into  that  continent. 

1  It  must  be  remembered  that  we  are  here  dealing  with  the  mammalian 
evidence  alone.  In  regard  to  the  molluscan  Mr  A.  H.  Cooke  (Conchologist, 
1893,  p.  131)  states  that  this  region  possesses  sufficient  individuality  from  that 
of  the  mainland  to  entitle  it  to  separation.  The  Helicida  are  peculiar,  not 
being  found  in  the  Mascarenes,  Seychelles,  or  Comoros.  They  seem  to  be 
related  to  certain  Cingalese  and  Australian  types.  Upwards  of  fifty-four  species 
of  Cyclostoma  are  known,  distributed  over  Madagascar,  the  Comoros,  Seychel- 
les, Mauritius,  and  Bourbon.  The  African  Bulimi  are  represented  by  two 
species,  but  Achatina  (so  common  there)  is  scarce;  and  groups  of  Bulimi  are 
peculiar.  A  single  species  of  the  genus  Caliella  is  identified  with  an  Indian 
form ;  and  unmistakable  indications  of  Oriental  affinities  are  afforded  by  the 
freshwater  molluscs.  There  are  two  species  of  Paludomus,  Bithynia  occurs, 
and  while  several  of  the  Melania  are  of  a  type  common  in  the  Indo- Malayan 
countries,  the  Melanatria,  which  are  peculiar  to  Madagascar,  have  their  nearest 
allies  in  Ceylon  or  India.  Although  not  a  single  African  freshwater  bivalve 
has  yet  been  recorded  from  Madagascar,  yet  several  Ethiopian  genera  of 
gastropods  occur  there,  and,  in  common  with  the  land-molluscs,  indicate  a 
former  connection  between  Madagascar  and  Africa,  and  this,  in  Mr  Cooke's 
opinion,  occurred  at  an  immeasurably  remote  epoch. 


224  THE   MALAGASY   REGION.  [CHAP. 

been  found  in  the  island.  After  remarking  that  bush-pigs  are 
stated  to  be  more  aquatic  in  their  habits  than  ordinary  swine,  Dr 
Blanford1  asks  "how  far  could  Potamochxrus  swim?  Surely  it 
is  not  likely  that  it  could  cross  the  Straits  of  Dover.  I  think  we 
are  justified  in  assuming  about  ten  miles  as  a  probable  limit  of  its 
power  of  crossing  the  sea,  but,  to  be  safe,  let  us  suppose  double  as 
much.  Then,  in  Pliocene  or  Plistocene  times,  quite  as  probably 
the  latter  as  the  former,  when  Potamochczmis  reached  South  Africa, 
Madagascar  was  separated  by  a  channel  not  more  than  twenty 
miles  broad.  The  conclusion  is  inevitable,  that  nearly  the  whole 
depression  of  upwards  of  a  thousand  fathoms  is  of  Pliocene  or 
Post-pliocene  date.  Of  course  it  must  not  be  assumed  that  this 
date  is  proved.  What  we  may  consider,  however,  as  beyond  any 
doubt  is  that  the  depression  cannot  be  older  than  the  Middle 
Tertiary."  This  view  may  be  taken  as  practically  identical  with 
the  one  here  advanced,  namely  that  Africa  and  Madagascar  were 
united  till  the  period  of  the  upper  Oligocene  or  Miocene. 

With  the  exception  of  the  fruit-bats  and  Daubenton's  civet, 
which,  as  already  mentioned,  is  more  nearly  allied  to  the  Oriental 
rasse  than  to  the  Ethiopian  Viverridce,  the  Malagasy  mammals  do 
not  exhibit  any  well-marked  alliance  with  those  of  India.  But  the 
case  is  different  with  the  birds,  molluscs,  and  certain  other  groups  ; 
while  we  have  no  evidence  that  giant  land-tortoises  ever  inhabited 
the  African  mainland,  although  an  extinct  species  is  known  from 
the  Indian  Pliocene. 

Basing  his  conclusion  on  evidence  drawn  from  several  sources, 
Dr  Blanford,  in  the  communication  last  cited,  is  of  opinion  that 
there  was  formerly  a  direct  land-connection  between  India  and 
South  Africa,  and  that  this  connection  "included  the  Archaean 
masses  of  the  Seychelles  and  Madagascar,  that  it  continued 
throughout  upper  Cretaceous  times,  and  was  broken  up  into 
islands  at  an  early  Tertiary  date.  Great  depression  must  have 
taken  place,  and  the  last  remnants  of  the  islands  are  now  doubt- 
less marked  by  the  coral  atolls  of  the  Laccadives,  Maldives,  and 
Chagos,  and  by  the  Saya  de  Malha  bank.  It  is  immaterial 
whether  Bourbon,  Mauritius,  and  Rodriguez  ever  formed  part  of 

1  Appendix,  No.  8,  p.  88. 


VI.]  RELATION    TO   AFRICA.  225" 

the  Mascarene  land  or  not."  It  is  added  that  if  future  soundings 
should  indicate  the  absence  of  a  bank  extending  the  whole  way 
from  India  to  Africa,  it  may  be  a  question  whether  the  whole  of 
the  ocean-bed  between  those  two  countries  has  not  sunk  to  its 
present  depth  since  the  Cretaceous  era1. 

This  presumed  connection  satisfactorily  explains  much  in 
regard  to  the  distribution  of  the  molluscs.  It  is,  however,  certain 
that  fruit-bats  did  not  exist  in  the  early  Tertiary,  and  the  Pteropus 
must  accordingly  have  made  the  journey  across  the  sea  from 
India,  aided  by  what  remained  of  the  chain  of  islands,  which  may 
have  been  more  extensive  during  the  Pliocene.  The  same 
explanation  also  holds  good  with  regard  to  most  of  the  Oriental 
types  of  birds.  The  case  of  the  land-tortoises  is,  however,  more 
difficult.  Nearly  allied  forms  have  been  found  in  Mauritius, 
Rodriguez,  Madagascar,  and  Aldabra;  and  since  this  group  is 
unknown,  even  in  Europe,  before  the  Oligocene,  it  is  evident  that 
they  could  not  have  travelled  from  India  by  means  of  the  con- 
necting land-bridge,  which  is  considered  to  have  been  broken  up 
at  the  commencement  of  the  Tertiary  epoch.  This  being  so,  the 
probability  is  that  they  originally  came  from  Africa ;  but  whether 
they  entered  Madagascar  with  the  ancestral  lemurs,  or  whether 
they,  or  their  eggs,  were  transported  across  the  channel  when 
narrower  than  at  present,  there  is  no  evidence  to  show.  Be  this 
as  it  may,  it  is  probable  that  they  reached  Rodriguez  and  Mauritius 
across  the  intervening  sea,  since  even  if  these  islands  ever  joined 
Madagascar,  such  union  must  apparently  have  been  at  a  date 
anterior  to  the  existence  of  true  tortoises.  That  none  of  these 
tortoises  could  have  been  transported  by  sea  from  India  is  proved 
by  an  observation  of  Dr  Blanford  to  the  effect  that  on  this  line  the 
currents  invariably  set  from  the  Seychelles  to  India.  It  may  be 
added  that  some  writers  have  considered  it  probable  that  the  giant 
tortoises  of  the  Malagasy  region,  like  those  of  the  Galapagos 
Islands,  attained  their  large  dimensions  after  they  had  reached 
the  islands  they  respectively  inhabit.  The  existence  of  gigantic 

1  Neumayr  (Erdgeschichte,  2nd  ed.  vol.  n.  p.  262,  1895)  considers  that  when 
India  was  connected  with  Madagascar  during  the  Jurassic  era,  only  the  southern 
extremity  of  that  island  was  joined  to  South  Africa. 

L.  15 


226  THE   MALAGASY   REGION.  [CHAP.  VI. 

species   on   nearly  all  the  great  continents    during  the  Tertiary 
epoch  seems,  however,  an  insuperable  objection  to  this  view. 

In  the  absence  of  any  evidence  as  to  the  Tertiary  vertebrate 
palaeontology  of  eastern  Africa,  I  have  no  suggestion  to  offer  as  to 
the  origin  of  the  gigantic  birds  of  the  genus  SEpyornis,  which 
during  the  late  Plistocene  or  recent  epoch  formed  such  a  marked 
feature  in  the  Malagasy  avifauna. 

NOTE. — The  Author  has  reason  to  believe  that  several  new 
Malagasy  mammals  have  been  discovered  by  Dr  C.  I.  Forsyth-Major; 
but  as  no  description  of  these  had  appeared  when  this  chapter  was 
passed  for  press,  they  could  not  be  noticed. 


CHAPTER   VII. 

THE    ETHIOPIAN    REGION. 

Extent — Characteristics  of  the  Mammalian  Fauna — Birds — Past  History  of 
Ethiopia — Subregions. 

IT  would  be  difficult  to  find  a  much  greater  contrast  to  the 
mammalian  fauna  of  Madagascar  than  is  presented  by  that  of 
Africa  south  of  the  tropic  of  Cancer ;  the  one  area,  as  shown  in  the 
last  chapter,  being  characterised  by  the  number  of  lemurs,  together 
with  its  peculiar  Viverridce.  and  insectivores,  while  the  other  is 
distinguished  from  all  other  parts  of  the  world  by  the  extraordinary 
number  (both  as  regards  genera,  species,  and  individuals)  of  large 
ungulates  which  roamed  through  its  plains  and  forests  until  deci- 
mated or  exterminated  by  the  hand  of  man.  As  regards  the 
number  of  individuals  of  large  animals  inhabiting  equal  areas,  it  is 
quite  probable  that  at  the  date  when  the  bison  flourished  in  its 
millions  on  the  North  American  prairies,  the  balance  in  this 
respect  may  have  been  in  favour  of  the  New  World  ;  but  whereas 
the  prairies  had  but  a  single  species,  Ethiopian  Africa  was  popu- 
lated (for  it  is  unfortunately  necessary  to  write  in  the  past  tense) 
with  a  host  of  species  of  antelopes,  together  with  buffaloes,  giraffes, 
hippopotami,  zebras,  rhinoceroses,  and  elephants.  Such  a  fauna 
has  existed  during  the  recent  epoch  in  no  other  part  of  the  world, 
and  in  past  times  has  only  been  paralleled  by  the  lower  Pliocene 
fauna  of  southern  Europe  and  Asia,  although  even  this,  as  regards 
the  number  of  generic  and  specific  types  of  antelopes,  is  by  no 
means  its  equal. 

Separating  Madagascar  and  the  associated  islands  as  a  distinct 
division,  the  Ethiopian  region  may  be  taken  to  include  such 
portions  of  Africa  and  Arabia  as  lie  to  the  south  of  the  tropic  of 

15—2 


228  THE    ETHIOPIAN    REGION.  [CHAP. 


Cancer ;  northern  Africa,  as  it  did  in  the  Pliocene,  clearly  forming 
a  part  of  the  Holarctic  region.  The  greater  part  of 
the  Sahara,  as  well  as  the  northern  portion  of 
the  Nubian  desert,  although  included  in  the  Holarctic,  will  form 
a  kind  of  transition  zone  towards  that  region,  as  is  also  the  case 
with  Syria,  where  a  considerable  number  of  Ethiopian  types  of 
mammals  are  met  with,  while  western  Arabia  shows  a  decided 
approximation  to  the  Oriental  region,  as  is  well  exemplified  by 
the  occurrence  there  of  a  species  of  the  short-horned  goats  con- 
stituting the  genus  Hemitragus.  As  has  been  stated  in  an  earlier 
chapter,  the  Sahara  and  Nubian  deserts,  although  they  have 
apparently  never  been  submerged  since  the  Cretaceous  epoch,  seem 
always  to  have  formed  a  more  or  less  complete  barrier  to  the  passage 
of  the  mammals  of  Algeria  and  the  adjacent  countries  into  the 
Ethiopian  region ;  and  the  main  migration  from  the  north  and  east 
has  thus  taken  place  along  the  north-eastern  side  of  the  continent. 
With  the  exception  of  its  southern  extremity,  the  whole  of  this 
vast  area  lies  within  the  tropics.  As  regards  its  physical  features, 
Dr  Heilprin  writes  that  "  it  presents  several  well-marked  physical 
peculiarities.  In  the  first  place,  we  have  the  vast  expanse  of 
desert,  which  in  the  north  occupies  a  transverse  band  varying  in 
width  from  about  four  to  nearly  ten  degrees  of  latitude.  This  is 
succeeded  by  what  may  not  improperly  be  termed  the  open 
pasture-lands,  which  as  a  narrow  belt  bounds  the  Sahara  on  the 
south,  curves  southwards  at  about  the  position  of  Kordofan,  and 
occupies  the  greater  portion  of  the  continent  lying  east  of  the 
thirtieth  parallel  of  east  longitude  and  south  of  the  fifth  parallel  of 
south  latitude.  A  very  considerable  portion  of  this  pasture-tract 
forms  a  plateau  of  from  four  thousand  to  five  thousand  feet  eleva- 
tion. Included  within  it,  and  bounded  on  the  west  by  the  Atlantic 
Ocean,  is  the  region  of  the  great  equatorial  forests,  to  the  present 
day  a  terra  incognita  in  great  part  both  to  geographers  and 
naturalists.  That  portion  of  the  African  continent  lying  south  of  the 
tropic  of  Capricorn  differs  in  many  respects,  both  as  to  its  physical 
configuration  and  its  vegetable  products,  from  the  region  to  the 
northward,  and  is  characterised  by  a  vegetation  which  is  one  of 
the  richest  and  most  remarkable  on  the  globe.  With  this  marked 
peculiarity  in  its  vegetable  development  there  is  of  necessity  a 


VII.]  SUB-REGIONS.  229 

certain  amount  of  faunal  peculiarity  superadded  as  well,  but  this  is 
not  sufficiently  pronounced  to  permit  of  the  separation  of  this 
tract  from  the  tract  lying  immediately  to  the  north.  We  have  thus 
on  the  continent  three  strictly  defined  faunal  sub-regions  :  (i)  the 
pasture-lands  already  described,  constituting  the  East  Central 
African  sub-region,  through  whose  vast  expanse  there  is  manifest 
a  strong  identity  in  the  character  of  the  animal  products,  the  same 
or  very  closely  related  forms  being  in  many  instances  found  at  the 
extreme  points  of  this  sub-region  ;  (2)  the  forest-tract,  constituting 
the  West  African  sub-region,  whose  animal  products  naturally 
differ  very  essentially  from  those  of  the  last ;  and  (3)  the  desert  or 
Saharan  sub-region,  containing  a  comparatively  limited  fauna, 
which,  with  almost  insensible  gradations,  merges  into  the  fauna  of 
the  Mediterranean  tract.  To  the  same  division  belong  in  great 
measure  the  desert  tracts  of  Arabia,  or  that  portion  of  the 
peninsula  lying  to  the  south  of  the  tropic  of  Cancer." 

Although  Dr  Wallace  had  previously  divided  continental 
Ethiopia  into  an  East  African,  West  African,  and  South  African 
sub -region,  the  foregoing  arrangement  seems,  on  the  whole, 
preferable.  There  are,  however,  considerable  reasons  for  regard- 
ing Somaliland  as  a  sub-region  by  itself,  and  South  Arabia  should 
perhaps  constitute  another.  Although  the  precise  determination 
of  such  areas  does  not  come  within  the  province  of  this  work,  it  is 
most  important  to  notice  that  the  West  African  or  Equatorial 
forest  tract  continues  right  across  the  continent  as  far  eastwards  as 
the  Congo- Nile  watershed,  that  is  to  say,  close  up  to  Wadelai, 
where  all  traces  of  the  West  African  fauna  are  suddenly  lost.  On 
this  point  Mr  O.  Thomas1  writes  that  "the  abruptness  with  which 
the  change  of  fauna  occurs  on  the  watershed  is,  considering  the 
insignificant  nature  of  the  physical  barriers,  very  remarkable,  and 
almost  unequalled  in  the  distribution  of  the  mammals  of  any  part 
of  the  world.  The  reason  of  the  change  is,  however,  clear  enough, 
being  not  the  occurrence  of  such  barriers  to  migration  as  moun- 
tains or  rivers,  but  the  abrupt  ending  of  the  great  West  African 
forest,  which,  as  we  know  from  the  travels  of  Schweinfurth  and 
ethers,  extends  quite  into  this  region,  but  abruptly  ceases  before 
the  slopes  of  the  upper  Nile  basin  are  reached." 
1  Proc.  Zool.  Sec.  1888,  p.  17. 


230  THE   ETHIOPIAN    REGION.  [CHAP. 

Before  considering  the  leading  characteristics  of  the  Ethiopian 

mammal  fauna  and  its  relations   to  that  of  other 

characteris-     regions,   both  in  the  present  and    the   past,   it  is 

tics  of  Mam-  .  & .      . 

maiian  Fauna,     desirable  to  make  reference  to  certain  deficiencies, 
which  are  very  difficult,  if  not  impossible  to  explain 
adequately  with  our  present  knowledge. 

Although  deer  (Cervus),  typical  pigs  (the  genus  Sus  in  its 
restricted  sense),  and  bears  are  met  with  in  northern  Africa,  no 
member  of  any  one  of  these  genera  with  the  single  exception  of 
a  pig  (Sus  sennaarensis)  from  the  Sennaar  district  of  Upper  Nubia, 
inhabits  Ethiopia.  Even  the  entire  family  of  the  Cervtdce  is 
unrepresented.  These  deficiencies  form  a  most  marked  contrast 
between  the  Ethiopian  region  on  the  one  hand,  and  both  the 
Oriental  and  the  Holarctic  on  the  other.  Almost  equally  con- 
spicuous is  the  absence  of  goats  and  sheep  ;  the  only  exceptions 
being  the  occurrence  of  a  species  of  Capra  in  the  highlands  of 
Abyssinia,  and  one  of  Hemitragus  in  Oman,  in  south-eastern 
Arabia.  The  absence  of  sheep  and  goats  is,  however,  by  no  means 
so  remarkable  as  that  of  the  other  groups  above  mentioned,  since 
the  former  are  exclusively  mountain  animals,  and  probably  need 
some  general  lowering  of  the  temperature  to  enable  them  to  pass 
from  one  chain  to  another,  and  of  the  existence  of  such  cold 
period  there  seems  no  evidence  in  Ethiopian  Africa.  A  somewhat 
similar  explanation  will  probably  apply  to  the  total  absence  of 
marmots  (Arctomys\  susliks  (Spermophilus],  chipmunks  (Tamias), 
beavers  (Castoridce],  voles  (Microtince),  and  picas  (Lagomys),  since 
all  these  are  inhabitants  of  elevated  or  northern  areas.  More  diffi- 
cult to  explain  is  the  absence  of  all  shrews  (Soricidce),  with  the 
exception  of  one  genus  peculiar  to  the  region ;  but  the  deficiency 
of  moles  (TalpidcR)  may  perhaps  be  accounted  for  by  the  slow 
travelling  powers  of  these  animals,  which  did  not  allow  them 
time  to  pass  into  Ethiopia  during  the  (probably  short)  period 
when  its  connection  with  other  regions  was  of  such  a  nature  as  to 
permit  their  living  in  the  intermediate  lands.  Possibly  also  the 
absence  of  moles  from  peninsular  India  has  something  to  do  with 
this  deficiency.  In  this  connection  it  is  worth  remark  that  the 
place  held  in  the  Holarctic  region  by  moles  is  by  no  means 
unoccupied  in  the  Ethiopian,  both  the  golden  moles  ( Chrysochloris), 


VII.]  PRIMATES.  231 

and  the  so-called  Cape  mole  (Bathyergus),  with  its  allies,  having 
similar  subterranean  habits. 

Together  with  the  Oriental,  the  Ethiopian  region  shews  a 
marked  distinction  from  all  others  as  the  sole  habitat  of  the 
man-like  apes  (Simiidce).  The  Ethiopian  forms  comprise  the 
chimpanzees  (Anthropopithecus)  and  gorilla  (Gorilla),  both  of 
which  are  restricted  to  the  equatorial  forest-region,  where  the 
former  ranges  as  far  east  as  Uganda,  although  the  latter  has  a 
more  circumscribed  distribution.  The  occurrence  of  a  fossil 
chimpanzee  in  the  Indian  Pliocene  affords  the  most  convincing 
evidence  of  the  derivation  of  a  large  part  of  the  Ethiopian  fauna 
from  what  is  now  the  Oriental  region.  Among  the  ordinary 
monkeys  and  baboons  (Cercopithetida)  there  are  five  genera  con- 
fined to  this  region.  Of  these,  Colobus  differs  from  the  Oriental 
langurs  (Semnopithecus]  by  the  absence  or  rudimentary  condition  of 
the  thumb,  which  frequently  has  lost  all  trace  of  a  nail.  On  the 
other  hand,  the  large  genus  Cercopithecus^  which  is  most  fully 
represented  in  the  forest  region,  is  as  nearly  related  to  the  Oriental 
Macacus,  from  which  it  differs  in  the  less  prominent  muzzle,  and 
the  absence  of  a  projecting  heel,  or  hinder  lobe  to  the  last  lower 
molar  tooth.  This  heel  is,  however,  present  in  the  mangabeys,  or 
white-eyelid  monkeys  (Cercocebus),  all  of  which  are  exclusively 
confined  to  the  forest  tract.  Although  the  dog-faced  baboons 
(Papio1)  have  a  wider  distribution,  ranging  from  the  Cape  to 
Arabia,  some  of  the  largest  and  most  peculiar  forms,  such  as  the 
mandrill  and  drill,  are  confined  to  West  Africa.  This  genus  is  one 
of  those  common  to  the  Ethiopian  region  and  the  Indian  Pliocene. 
The  nearly-allied  gelada  baboons  (Theropithecus],  of  which  there 
are  two  representatives,  are,  on  the  other  hand,  exclusively  north- 
eastern types,  one  being  confined  to  Abyssinia.  Among  the 
lemuroids,  the  galagos  (Galago*) — which,  as  stated  in  the  last 
chapter,  belong  to  a  sub-family  which  attains  its  maximum  develop- 
ment in  Madagascar — extend  right  across  the  equatorial  portion  of 
the  continent,  descending  somewhat  on  the  east  coast,  where  they 
are  very  numerously  represented.  The  pottos  (Perodicticus],  which 
are  nearly  related  to  the  lorises  of  the  Oriental  region,  are,  how- 

1  Syn.  Cynocephahis.  *  Including  Otogale. 


232 


THE    ETHIOPIAN    REGION. 


[CHAP. 


ever,  exclusively  confined  to  West  Africa,  where  they  are  known 
by  two  species,  regarded  by  some  as  representing  as  many  genera. 

It  will  be  unnecessary  to  say  more  with  regard  to  the  Chiro- 
ptera  than  that  the  fruit-bats  (Pteropodidce)  are  represented  solely  by 
three  peculiar  genera  in  the  Ethiopian  region,  of  which  Epomo- 
phorus  has  the  greater  number  and  the  most  peculiar  of  its  species 
confined  to  the  western  forest  tract,  while  the  single  species  of 
Scotonycteris  is  solely  found  in  this  part  of  this  continent,  as  is 
also  the  case  with  that  of  Trygenycteris. 

Of  great  importance  from  a  distributional  point  of  view  are  the 
Ethiopian  Insectivora,  since  of  the  five  families  found  within  the 


FIG.  52.     AFRICAN  JUMPING-SHREW  (Macroscelides  tetradactyhis]. 

area  under  consideration,  two  are  almost  or  exclusively  confined 
to  it,  while  the  third  has  only  one  aberrant  representative  in  Mada- 
gascar, and  even  this  may  prove  entitled  to  constitute  a  family  by 
itself  when  its  structure  is  more  fully  known  than  is  at  present  the 


VII.]  INSECTIVORA.  233 

case.  It  is  further  noticeable  that  two  of  the  families  belong  to 
the  primitive  group  characterised  by  their  tritubercular  upper  molar 
teeth,  and  were  accordingly  in  all  probability  very  early  immigrants 
into  the  country.  The  first  family  almost  peculiar  to  the  region  is 
that  of  the  jumping-shrews  (Macroscelididce),  the  members  of  which 
are  easily  recognised  by  their  elongated  hind  limbs,  long  snout, 
and  leaping  habits.  While  the  typical  genus  Macroscelides  has 
representatives  throughout  the  region  and  also  extends  into 
Holarctic  Africa,  the  four  species  of  Rhynchocyon,  in  which  the 
legs  are  shorter  and  the  snout  longer,  are  restricted  to  East  Africa. 
To  the  latter  is  closely  allied  the  European  Oligocene  genus 
Pseudorhynchocyon,  and  it  would  thus  seem  that  the  family, 
although  unrepresented  in  Madagascar,  arrived  at  a  relatively 
early  date  in  Ethiopia.  On  the  other  hand,  since  the  hedgehogs 
•  are  represented  only  by  the  widely-spread  typical  genus  Erinaceus, 
together  with  a  West  African  species  which  has  been  separated  as 


FIG.  53.     WEST  AFRICAN  POTAMOGALE  (Potamogale  velox]. 

Proechinus,  it  is  probable  that  they  did  not  make  their  appearance 
on  the  scene  till  a  later  epoch.  The  Soricidce,  or  shrews,  are 
represented  in  Ethiopia  only  by  three  species  belonging  to  the 
peculiar  genus  Myosorex,  differing  from  all  the  other  genera  in 


234  THE    ETHIOPIAN    REGION.  [CHAP. 


which  the  teeth  are  white  by  the  absence  of  long  hairs  on  the  tail1. 
With  the  West  African  genus  Potamogale,  we  come  to  the  first  of 
the  two  families  with  tritubercular  upper  molars ;  the  present  one 
(Potamogalidce]  being  represented  elsewhere  only  by  the  Malagasy 
Microgale,  of  which,  as  already  said,  the  systematic  position  is 
doubtful.  The  potamogales,  which  attain  a  couple  of  feet  in 
length,  are  thoroughly  aquatic  in  their  habits,  swimming  by  the  aid 
of  the  highly  compressed  tail.  It  has  generally  been  considered 
that  there  is  only  a  single  species,  but  it  has  recently  been 
suggested  that  there  may  be  two.  The  family  is  probably  an 
ancient  one,  although  we  have  no  fossil  evidence  to  this  effect, 
Microgale,  even  if  it  belong  to  another  family,  indicating  that  the 
group  was  among  the  earlier  mammalian  colonists  of  Ethiopia. 
The  golden  moles  (Chrysochloridcz],  which  take  their  name  from 
the  brilliant  metallic  lustre  of  the  fur  in  the  majority  of  the  species, 
are  blind,  earless,  fossorial  insectivores,  having  the  middle  toe  of 
the  fore  foot  furnished  with  an  enormously  powerful  claw.  As  the 
moles  (Talpida]  form  a  group  nearly  related  to  the  shrews,  so  the 
golden  moles  are  equally  nearly  related  to  the  tenrecs  ( Centetidcz) 
of  Madagascar,  from  which  they  may  be  regarded  as  a  highly 
specialised  offshoot.  Accordingly,  it  may  be  taken  for  granted 
that  their  ancestors  obtained  an  entry  into  Ethiopia  with  the 
ancestral  lemurs.  It  is  not  improbable  that  the  prevalence  of 
higher  types  of  mammalian  life  has  been  the  cause  of  the  assump- 
tion of  mole-like  habits  in  the  Chrysochloridce  ;  the  tenrecs,  which 
live  in  an  island  where  the  competition  is  much  less  severe,  having 
retained  the  original  primitive  type.  The  golden  moles,  which 
may  all  be  included  in  the  single  genus  Chrysochloris,  are  mainly 
confined  to  South  Africa,  although  one  species  extends  on  the 
east  coast  as  far  north  as  Ugogo. 

Turning  to  the  Carnivora,  it  is  unnecessary  to  say  anything 
with  regard  to  the  Felida,  except  that  three  species  of  felts  are 
common  to  Ethiopia  and  India;  while  the  single  species  of 
hunting-leopard  (Cyncelurus]  is  likewise  found  in  both  countries, 
the  genus  being  apparently  also  represented  in  the  Indian  Pliocene. 
In  the  civet  family  ( Viverridce)  the  true  civets  ( Viverra]  and 

1  See  Dobson,  Proc.  Zool.  Soc.  1887,  p.  575. 


VII.]  CARNIVORA.  235 

mungooses  (Herpestes]  are  common  to  the  Ethiopian  and  Oriental 
regions ;  but  the  whole  group  attains  a  far  greater  development 
within  the  former  area  than  elsewhere.  Although  the  common 
genet  is  an  inhabitant  of  the  southern  portion  of  the  eastern 
Holarctic  region,  all  the  other  species  of  Genetta  are  Ethiopian. 
The  West  African  linsang  (Poland]  is  the  Ethiopian  representative 
of  the  beautiful  linsangs  (Linsangd)  of  the  eastern  portion  of  the 
Oriental  region  ;  the  distribution  of  this  group  being  a  well-marked 
instance  of  the  close  alliance  of  the  fauna  of  the  Malayan 
countries  to  that  of  West  Africa,  to  which  reference  will  again  be 
made.  The  Oriental  palm-civets  (Paradoxurus)  are  represented 
by  the  nearly-allied  Ethiopian  genus  Nandinia,  of  which  one 
species  is  West  African,  while  the  other  comes  from  Nyasaland. 
The  small-toothed  mungoose  (Helogale]  is  common  to  West  and 
East  Africa ;  and  the  allied  genus  Bdeogale  has  representatives 
on  both  sides  of  the  continent.  Two  other  peculiar  Ethiopian 
genera,  Cynictis  and  Rhynchogale,  have  each  but  a  single  species  :  the 
former  being  South  African  and  the  latter  East  African.  The 
cusimanses  (Crossarchus),  although  mainly  characteristic  of  the 
forest  tract,  have  one  representative  in  Abyssinia.  Lastly,  the 
meerkat,  the  sole  representative  of  the  genus  Suricata,  is  an 
exclusively  South  African  form.  A  peculiar  family  (Proteleidce)  is 
constituted  by  the  aard-wolf  (Proteles],  ranging  from  the  Cape  to 
Somaliland,  and  a  near  ally  of  the  hyaenas,  from  which  it  is 
distinguished  by  the  extremely  feeble  development  of  the  denti- 
tion. Both  the  spotted  hyaena  (Hyczna  crocuta)  and  the  brown 
hyaena  (H.  fused)  are  now  confined  to  Ethiopia,  but  the  former 
ranged  over  a  large  portion  of  Europe  as  well  as  southern  India 
during  the  Plistocene  epoch ;  and  as  all  the  three  living  species 
are  included  in  the  same  genus,  there  is  no  generic  type  in  this 
family  restricted  to  the  Ethiopian  region. 

In  the  Canidcz  wolves  are  absent,  but  the  jackals  are  repre- 
sented by  species  allied  to  Cam's  aureus,  which  occurs  in  North 
Africa;  wild  dogs  (sub-genus  Cyon)  are,  however,  wanting. 
Although  the  long-eared  foxes  or  fennecs,  such  as  Cants  chama, 
are  common  in  Ethiopia,  they  are  by  no  means  characteristic, 
since  they  range  into  North  Africa,  Syria,  Persia,  and  Afghanistan  ; 
being,  in  fact,  like  the  gazelles,  desert-haunting  forms.  There  are, 


236 


THE    ETHIOPIAN    REGION. 


[CHAP. 


however,  two  genera  of  the  family,  each  with  a  single  species,  now 
confined  to  this  region  ;  the  first  being  represented  by  the  Cape 
hunting-dog  {Lycaon  pictus],  which  is  a  large,  somewhat  hy?ena-like 
animal,  easily  recognised  by  its  spotted  coloration  and  long  bushy 
tail,  and  distinguished  from  the  other  genera  by  having  only  four 
toes  on  both  the  fore  and  hind  feet.  That  the  genus  is  of  northern 


FIG.  54.     CAPE  HUNTING-DOG  (Lycaon  pictits). 

origin  is  proved  by  the  occurrence  of  remains  of  an  extinct  species 
in  the  Glamorganshire  caves.  The  small  Lalande's  fox  (Otocyon 
megalotis)  of  South  Africa,  in  addition  to  the  enormous  size  of  its 
ears,  is  peculiar  in  having  four  pairs  of  molar  teeth  in  the  lower 
jaw,  and  either  three  or  four  in  the  upper.  Possibly  a  fossil  species 
from  the  Indian  Pliocene  may  be  an  allied  type. 

Passing  by  the  Ursidce.  and  Procyonidce  as  unrepresented  in 
this  region,  we  find  the  Mustelida  very  poorly  developed,  martens 
(Mustela)  being  absent,  and  true  weasels  very  scarce.  The  striped 
Gape  weasel  (Poedlogale]  constitutes,  however,  a  genus  by  itself; 
while  the  similarly  coloured  Cape  polecat  (fctonyx),  is  one  of  two 
representatives  of  a  small  genus,  the  second  of  which  ranges  from 


VII.] 


RODENTIA. 


237 


Sennaar  to  Egypt,  and  is  also  stated  to  occur  in  Asia  Minor.  A 
fossil  species  from  the  European  Miocene  may  perhaps  belong  to 
this  genus.  The  ratels  (Mellivora)  which  are  now  represented  by 
one  Ethiopian  and  a  second  Indian  species,  are  proved  to  be  com- 
paratively late  immigrants  from  the  north  into  this  region  by  the 
occurrence  of  a  fossil  species  in  the  lower  Pliocene  of  Northern 
India.  As  the  animal  described  under  the  name  of  Galeriscus  is 
at  present  known  only  by  a  skin,  it  is  not  even  certain  that  it 
belongs  to  the  Mustelidce.  at  all. 

Among  the  squirrel-like  rodents,  the  most  striking  feature  is 
the  absence  of  the  true  flying-squirrels  and  their  replacement  by  a 


FIG.  55.     FULGENT  AFRICAN  FLYING-SQUIRREL  (Anomalurus  fulgens). 

distinct  family  (Anomaluridce.},  characterised,  among  other  features, 
by  the  presence  of  scales  on  the  under  surface  of  the  root  of  the 
tail.  Mainly  characteristic  of  the  forest  area,  the  typical  genus 
Anomalurus  has,  nevertheless,  representatives  on  the  eastern  side 


238  THE    ETHIOPIAN    REGION.  [CHAP. 


of  the  continent;  although  the  mouse-like  long-tailed  flying- squirrel 
(Idiurus)  is  exclusively  West  African.  While  true  squirrels  (Sci- 
urus]  are  common  to  this  and  the  other  regions  of  Arctogaea,  the 
pigmy  squirrels  constituting  the  genus  Nannosciurus  are  repre- 
sented by  one  species  (N.  minutus)  in  West  Africa,  while  all  the 
others  are  Malayan.  The  spiny  squirrels  of  the  genus  Xerus  are 
now,  on  the  other  hand,  exclusively  Ethiopian,  although  their 
northern  origin  is  proclaimed  by  the  occurrence  of  an  extinct 
species  in  the  French  Miocene.  Dormice  (Myoxidce)  are  exceed- 
ingly abundant  in  Ethiopia,  and  if  it  is  considered  desirable  to 
split  up  the  family  into  more  than  two  genera,  the  genus  Graphi- 
urus,  characterised  by  the  short,  cylindrical,  and  tufted  tail,  and 
the  simple  structure  of  the  molar  teeth,  will  be  peculiar  to  this 
region,  as  will  also  be  the  single  West  African  form  described  as 
Claviglis.  Dormice,  as  mentioned  earlier,  date  in  Europe  from 
the  lower  Oligocene,  and  therefore  they  might  well  have  entered 
Ethiopia  with  the  ancestral  lemuroids,  although,  so  far  as  it  goes, 
their  absence  from  Madagascar  is  against  this  view.  In  the 
mouse-family  (Muridce)  five  sub-families  are  met  with  in  Ethiopia, 
two  of  which  are  peculiar  to  the  region.  Curiously  enough,  the 
cricetine  sub-family,  which  is  the  oldest  of  all,  and  the  only  one 
met  with  in  Madagascar,  is  represented  only  by  a  single  highly 
specialised  genus  (Trilophomys).  The  inference  from  this  would 
seem  to  be  that  the  ancestral  cricetines  and  lemuroids  entered 
Ethiopia  together,  whence  some  migrated  to  Madagascar ;  the 
former  group,  with  the  exception  of  the  one  peculiar  genus,  having 
completely  died  out  on  the  continent,  where  the  remaining 
murines  are  more  recent  immigrants  from  the  north.  In  this 
family  the  Eastern  Arctogaeic  group  of  the  gerbils  (Gerbillince) 
includes  six  genera,  out  of  which  no  less  than  five,  namely 
Pachyuromys,  Mystromys,  Otomys,  Dasymys,  and  Malacomys,  are 
exclusively  Ethiopian,  the  last  being  West  African.  The  elongated 
hind  limbs  and  the  transverse  laminae  of  the  molars  readily  serve 
to  distinguish  the  gerbils  from  the  exclusively  Ethiopian  sub- 
family Dendromyin&i  in  which  the  molars  are  rooted  and  tuber- 
culated  and  the  ears  remarkably  hairy.  The  typical  genus 
Dendromys  includes  two  dormouse-like  forms,  one  from  South, 
and  the  other  from  East  Africa;  the  other  genera  being  Lima- 


VII.]  RODENTIA.  239 

comys,  Steatomys,  and  Lophuromys,  the  last  having  the  fur  replaced 
by  fine  flattened  bristles.  Although  there  is  no  palaeontological 
history  of  either  of  the  preceding  sub-families,  the  Cricetince  have 
already  been  shown  to  date  from  the  lower  Oligocene  of  Europe. 
Their  sole  Ethiopian  representative  is  the  single  species  of  the 
genus  Trilophomys^,  from  Upper  Nubia  and  the  Red  Sea  littoral 
in  the  neighbourhood  of  Suakin,  and  perhaps  ranging  into  southern 
Arabia.  The  crested  rat,  as  the  creature  is  called,  takes  its  name 
from  the  prominent  crest  of  stiff  hair  running  down  the  back ; 
while  it  is  specially  characterised  by  the  roofing  over  of  the  whole 
upper  surface  of  the  skull  with  bone,  on  which  account  it  has 
(quite  unnecessarily)  been  made  the  type  of  a  family  by  itself. 
Perhaps,  however,  the  most  interesting  member  of  the  family 
inhabiting  Ethiopia  is  a  species  of  mouse  known  as  Deomys,  repre- 
senting both  a  genus  and  sub-family  by  itself,  and  characterised  by 
having  its  upper  molars  intermediate  between  those  of  the  crice- 
tines  and  murines.  Doubtless,  therefore,  this  rodent  is  a  somewhat 
modified  descendant  of  the  true  cricetines  which  entered  Africa 
while  it  was  still  united  to  Madagascar;  its  habitat  being  that 
refuge  for  ancient  types,  the  lower  Congo  valley.  The  two  other 
generic  representatives  of  the  family,  Cricetomys  and  Saccostomus, 
although  resembling  the  hamsters  in  the  presence  of  cheek- 
pouches,  have  molars  like  other  Murina,  and  are  accordingly 
referred  to  that  sub-family.  While  there  are  two  species  of  the 
second  genus,  there  is  but  one  of  the  first.  By  some  zoologists 
the  striped  mice,  as  typically  represented  by  the  Barbary  mouse, 
are  separated  from  Mus  to  form  a  genus  Arvicanthis* ,  which  is 
mainly  Ethiopian. 

The  small  family  of  the  Spalacidce,  or  burrowing-rats,  has  four 
genera  out  of  six  exclusively  Ethiopian,  while  a  fifth  (Rhizomys\ 
which  is  mainly  Oriental,  enters  Abyssinia.  Of  the  four  Ethiopian 
genera,  which  constitute  a  sub-family  by  themselves,  Bathyergus 
includes  only  the  great  Cape  mole-rat  of  South  Africa;  and 
Myoscalops  has  also  but  a  single  species,  although  there  are  several 
of  Georhychus.  Closely  allied  to  the  latter  are  two  tiny  little 
burrowing  and  nearly  naked  creatures  (Heterocephalus)  from 

1  Syn.  Lophiomys.  '*  Syn.  Isomys. 


240  THE   ETHIOPIAN    REGION.  [CHAP. 

Somaliland,  which  may  be  regarded  as  degraded  descendants  from 
that  type.  The  family  is  unknown  either  in  Madagascar  or  in  a 
fossil  state  in  Europe,  although  an  extinct  species  of  Rhizomys 
occurs  in  the  Indian  Pliocene,  and  it  is,  therefore,  in  all  proba- 
bility a  comparatively  late  immigrant  into  Ethiopia.  In  addition 
to  one  genus  ranging  from  Nubia  to  Siberia,  the  jerboa-family 
(DipodidcE)  has  one  peculiar  Ethiopian  genus,  represented  solely 
by  the  Cape  jumping-hare  (Pedetes) ; — a  form  so  different  from  all 
the  others  that  it  must  constitute  a  sub-family  apart.  With  this 
genus  we  leave  the  mouse-like,  and  come  to  the  porcupine-like 
group  of  the  rodent  order,  in  which  the  family  Octodontidce.  has 
nearly  all  of  its  representatives  which  are  not  Neogaeic  confined  to 
this  region,  although  the  gundi  (Ctenodactylus)  of  North  Africa,  in 
the  neighbourhood  of  Tripoli,  is  south  Holarctic.  This  genus  and 
an  allied  type  (Pectinator)  from  Somaliland,  together  with  the  next 
form,  alone  represent  a  sub-family  typically  characterised  by  the 
presence  of  a  horny  comb-like  appendage  and  stiff  bristles  on  each 
of  the  hind  feet ;  Pectinator  thus  being  Ethiopian.  The  South 
African  rock-rat  (Petromys),  although  now  included  in  the  same 
group,  approximates  to  a  sub-family  of  which  all  the  members  are 
South  American ;  the  resemblance  between  one  of  the  latter  and 
the  Ethiopian  species  being  curiously  close.  The  two  species  of 
cane-rat  (Triaulacodus1)  constitute  the  sole  African  representatives 
of  a  sub-family  containing  a  large  number  of  Neogseic  genera: 
Probably,  as  there  has  already  been  occasion  to  remark,  both  the 
Neogaeic  and  Ethiopian  representatives  of  this  family  trace  their 
origin  to  the  extinct  Theridomyidce  of  the  Oligocene  of  the  Hoi- 
arctic  region  or  some  nearly  allied  forms ;  and  as  certain  forms 
occur  in  the  Santa  Cruz  beds  of  Patagonia,  it  is  probable  that  the 
migration  into  Ethiopia  was  at  least  as  early  as  that  of  the  early 
lemuroids,  the  extinct  Pellegrinia  of  the  Sicilian  Pliocene  being 
the  last  survivor  of  the  group  in  the  northern  half  of  the  Old 
World2.  Although  there  is  no  generic  type  of  porcupine  (Hystri- 

1  This  name  is  proposed  to  replace  Aulacodus,  Temm.  (1827),  which  was 
preoccupied  in  1822  by  Eschscholtz  for  a  genus  of  Coleoptera. 

2  There  is  difficulty  in  this  respect  on  account  of  the  absence  of  hystrico- 
morphous  rodents  from  Madagascar;  but  perhaps  the  early  forms  entered  the 
west  side  of  the  continent,  while  the  lemurs  travelled  in  along  the  east. 


VII.]  UNGULATES.  241 

cidai)  peculiar  to  Ethiopia,  it  is  not  improbable  from  the  wide  dis- 
tribution and  antiquity  of  the  group,  that  these  rodents  also  entered 
tropical  Africa  at  a  comparatively  early  epoch.  The  deductions 
drawn  from  these  rodents  as  to  a  connection  between  Africa  and 
South  America  have  been  mentioned  in  Chapter  III. 

Among  all  the  striking  features  of  the  mammalian  fauna  of 
Ethiopian  Africa,  none  is  more  remarkable  than  the  enormous 
preponderance  of  ungulates,  many  of  which  are  of  great  corporeal 
bulk.  Of  these  a  large  number  of  genera  and  two  families  are 
absolutely  peculiar  to  this  region.  As  may  be  gathered  both  from 
their  absence  at  the  present  day  in  Madagascar,  and  the  late 
epoch  at  which  their  remains  are  found  in  the  Tertiaries  of  the 
Holarctic  and  Oriental  regions,  all  these  creatures  have  reached 
the  Ethiopian  region  but  recently.  The  Hippopotamida  is  one 
of  the  two  families  now  practically  peculiar  to  the  region,  the 
common  species  {Hippopotamus  amphibius)  having  ranged  over  a 
considerable  portion  of  Europe  during  the  Plistocene  and  upper 
Pliocene  ages,  while  even  in  the  beginning  of  this  century  it 


FIG.  56.     HEAD  OF  WART  HOG  {Phacochtxrus  cethiopicus). 

frequented  lower  Egypt.     The   pigmy  hippopotamus   (H.  liberi- 

ensis)  of  western  Africa,  which  is  referred  by  many  writers  to  a 

genus  apart,  and  more  resembles  the  pigs  in  its  mode  of  life, 

I,  1 6 


242  THE    ETHIOPIAN    REGION.  [CHAP. 


appears  to  be  more  nearly  allied  to  a  small  species  from  the 
Sicilian  and  Maltese  Pliocene.  As  already  mentioned,  with  the 
exception  of  a  single  species  (Sus  sennaariensis],  true  pigs  are 
unknown  in  Ethiopia,  their  place  being  taken  by  the  two  species 
of  bush-pigs,  forming  the  potamochoerine  group  of  the  same  genus, 
and  distinguished  by  the  simpler  structure  of  the  molar  teeth,  as 
well  as  by  the  tendency  of  the  front  premolars  to  fall  out  in  the 
adult.  The  reason  for  the  occurrence  of  a  third  species  of  the 
group  in  Madagascar  has  been  already  sufficiently  discussed1. 
Still  more  distinctive  of  the  region  are  the  hideous  wart-hogs 
(Phacocharus\  specially  characterised  by  the  facial  warts  from 
which  they  take  their  name,  the  huge  tusks,  and  the  great  com- 
plexity of  the  last  molar  tooth  in  each  jaw ;  the  tusks  and  these 
molars  being  frequently  the  only  teeth  remaining  in  aged  animals. 
It  is,  however,  very  noteworthy  that  certain  extinct  species  of  Sus 
from  the  Pliocene  of  India  and  Algeria  have  their  last  molars  of  a 
type  which  could  easily  be  developed  into  those  of  the  wart-hogs ; 
and  it  would  accordingly  seem  that  the  latter  are  comparatively 
recent  descendants  of  ordinary  pigs.  Although  wild  camels  are 
unknown  in  the  region,  the  Tragulida  are  represented  in  West 
Africa  by  the  water-chevrotain,  which  is  now  the  only  existing 
species  of  the  genus  Dorcatherium,  although  fossil  forms  occur  in 
the  Pliocene  of  India  and  the  European  Miocene ;  the  ancestral 
forms  having  probably  entered  the  region  from  India.  The  second 
ungulate  family  now  confined  to  Ethiopia  is  the  Giraffidce,  of 
which  there  appears  to  be  only  a  single  living  species,  although  the 
North  African  form  shows  a  decided  difference  in  coloration  from 
its  southern  brother.  The  occurrence  of  species  belonging  to  the 
existing  genus  Giraffa  in  the  lower  Pliocene  of  Greece,  Persia, 
India,  and  China,  shows  that  giraffes  came  into  Africa  with  the 
other  ruminants  ;  the  African  species  being  very  probably  the 
direct  descendant  of  the  extinct  Indian  one. 

Abounding  as  it  does  in  ungulates  in  general,  Ethiopian  Africa 
is  the  especial  home  of  the  antelope  group,  which  here  takes 
the  place  of  the  sheep  and  goats  so  characteristic  of  the  elevated 
districts  of  the  eastern  half  of  the  Holarctic  region.  Regarding 

1  Supra,  p.  223. 


VII.] 


UNGULATES. 


243 


their  distribution  in  the  Ethiopian  region,  Dr  Sclater  writes1  that 
although  "  antelopes  are  to  be  met  with  in  every  part  of  Africa, 
they  are  most  numerous  where  the  country  is  comparatively  open, 
and  where  there  are  grassy  plains  interspersed  with  sheltering 
bushes.  South  of  the  tropic  of  Capricorn  this  condition  generally 
prevails,  and  throughout  the  Cape  Colony  and  its  adjoining  terri- 
tory they  are — or,  at  all  events,  before  the  advent  of  a  European 


FIG.   57.     WATER-CHEVROTAIN  (Dorcatherium  aqiiaticum}. 

population  were, — everywhere  abundant.  The  early  settlers  at  the 
Cape  describe  antelopes  as  to  be  met  with  in  herds  of  thousands 
on  the  veldt,  and  in  parts  of  Africa  where  the  white  man  and  his 
destructive  firearms  have  not  yet  penetrated  a  similar  condition 
prevails  even  at  the  present  day.  When  we  advance  further  north 
and  meet  with  the  dense  forests  of  the  Niger  and  Congo  basins, 
we  find  the  mass  of  antelopes  holding  rather  to  the  more  open 
lands  on  the  eastern  coast,  throughout  which  they  are  to  be  met 


Natural  Science,  vol.  I.  p.  -255  (1892). 


1 6— 2 


244  THE    ETHIOPIAN    REGION.  [CHAP. 

with  in  great  abundance  up  to  Cape  Guardafui.  The  vast  plains 
traversed  by  the  Upper  Nile  and  its  tributaries  are  likewise  well 
stocked  with  antelope  life  ;  but  in  the  great  Sahara  only  some  of 
the  more  desert-loving  forms  are  to  be  found.  In  Senegambia 
again,  and  in  the  more  open  districts  on  the  West  Coast,  many 
forms  of  antelopes  occur,  but  they  cannot  rival  the  numbers  and 
varieties  of  those  of  Eastern  and  Southern  Africa."  Although 
most  of  the  genera  of  Ethiopian  antelopes  are  peculiar  to  the 
continent,  a  few  desert-haunting  types  range  into  southern  Arabia, 
and  hence  northwards  into  Syria,  where  they  enter  the  Holarctic 
region.  Many  of  the  groups  have  extinct  representatives  in  the 
lower  Pliocene  of  southern  Europe  and  India,  and  since  existing 
Ethiopian  genera  are  more  common  in  the  latter  than  in  the 
former  area,  it  seems  probable  that  the  great  migration  into 
Ethiopian  Africa  has  taken  place  from  the  east,  by  way  of  Syria  or 
Arabia.  As  such  extinct  genera  or  species  have  been  already 
noticed1,  it  will  suffice  to  take  a  very  brief  survey  of  the  genera 
now  mainly  or  exclusively  confined  to  Ethiopian  Africa. 

The  first  section  includes  the  hartebeests  and  their  allies  the 
bontebok  and  blesbok,  all  of  which  may  well  be  included  in  the 
genus  Bubalis,  although  the  two  latter  are  often  separated  as 
Damaliscus.  One  species  of  the  typical  group  ranges  into  Syria, 
while  a  second  is  an  inhabitant  of  Tunis.  To  the  same  section 
also  belong  the  wildebeests,  or  gnus  ( Connochcetes).  On  the  other 
hand  the  numerous  species  of  duikerboks  (Cephalophus)  consti- 
tute, so  far  as  Africa  is  concerned,  a  section  to  themselves.  They 
are,  however,  allied  to  the  Indian  four-horned  antelope  (Tetraceros\ 
and  it  is  not  improbable  that  they  are  represented  in  the  Pliocene 
of  the  Siwalik  Hills.  While  many  species  of  the  genus  are  found 
in  East  and  South  Africa,  the  largest  kinds  are  confined  to  the 
forest-districts  of  the  West  Coast.  The  small  African  antelopes 
classed  by  Sir  V.  Brooke  in  the  Ceruicaprince  and  included  in 
the  genera  Neotragus  and  Nanotragus  are  now  referred  by  Messrs 
Sclater  and  Thomas  to  a  section  apart,  under  the  name  of 
Nanotragin<z,  and  are  classed  in  six  genera.  Of  these,  Madoqua  ~, 
with  six  species,  includes  Salt's  antelope  (M.  saltiand)\  Nanotragus 

1  Supra,  pp.  197 — 206.  2  Syn.  Neotragtis. 


VII.]  UNGULATES.  245 

is  represented  only  by  the  minute  royal  antelope  (JV.  pygmaus} 
of  Guinea;  Nesotragus  is  typified  by  the  Zanzibar  steinbok 
(JV.  moschatus] ;  the  true  steinbok  (Raphiceros  campestris'}  forms 
the  fourth  genus ;  the  oribi  of  South  Africa  ( Oribia  scoparia)  is 
still  more  distinct;  while  the  well-known  klipspringer  (Oreotragus 
saltator},  which  ranges  from  the  Cape  to  Abyssinia,  differs  from 
all  the  rest  by  its  coarse  brittle  fur. 

The  Cervicaprince,  include  larger  forms.  Foremost  among 
these  is  the  South  African  rheebok  (Pelea) ;  while  the  water-buck 
and  its  allies  (Cobtts)  are  some  of  the  largest  of  all  antelopes. 
The  last  representative  of  this  section  (Cervicaprd)  is  typified  by 
the  South  African  rietbok,  but  there  are  also  other  species  in  West 
and  East  Africa.  The  fine  South  African  antelope  known  as  the 
pala  (sEpyceros],  together  with  an  allied  species  from  the  West 
Coast,  form  the  first  representatives  of  another  section.  Here 
belong  the  true  gazelles  (Gazella),  more  characteristic  of  the  desert 
tracts  of  the  eastern  half  of  the  Holarctic  region,  although  repre- 
sented in  South  Africa  by  the  somewhat  aberrant  springbok,  as 
well  as  by  several  more  typical  species  on  the  East  Coast.  Clarke's 
gazelle  (Ammodorcas)  of  Somaliland  is,  however,  the  sole  species 
of  an  exclusively  Ethiopian  genus  ;  and  the  same  is  the  case  with 
Waller's  gazelle  (Lithocranius\  of  which  the  range  extends  on  the 
East  Coast  from  Somaliland  to  Kilimanjaro.  The  single  species 
of  the  East  African  genus  Dorcatragus  seems  to  be  an  aberrant 
gazelle,  with  the  trunk-like  muzzle  of  Madoqua,  but  retaining 
the  small  gland-pits  of  the  type.  Yet  another  section  of 
antelopes  is  typified  by  the  beautiful  sable  antelope  and  its 
allies  (Hippotragus),  in  which  the  horns  sweep  backwards  in  a 
graceful  curve,  and  are  ringed  nearly  to  their  tips.  This  genus  is 
exclusively  Ethiopian,  but  in  the  one  typified  by  the  gemsbok 
(Oryx),  and  characterised  by  the  long  and  slender  horns  being 
either  straight  or  but  slightly  curved,  and  ringed  only  at  the  base, 
the  range  includes  all  the  desert  regions  of  Africa,  Arabia,  and 
Syria,  although  the  majority  of  the  species  are  Ethiopian.  To  the 
same  section  belongs  the  addax  antelope  (Addax),  but  although 
this  animal  occurs  as  far  south  as  latitude  18°  N.,  it  is  mainly  an 
inhabitant  of  the  deserts  of  North  Africa,  Arabia,  and  Syria.  The 
last  section  includes  some  of  the  largest  and  at  the  same  time  the 


246 


THE   ETHIOPIAN    REGION. 


[CHAP. 


most  beautiful  of  all  antelopes,  the  three  Ethiopian  genera  being 
all  characterised  by  the  more  or  less  strongly-marked  spiral  twisting 
of  their  horns,  and  the  short  crowns  of  their  molar  teeth  ;  the  last 
feature  distinguishing  them  sharply  from  the  sable  antelope  and 
gemsbok  group.  The  first  of  the  genera  in  question  includes  the 


FIG.  58.      HEAD  OF  GEMSBOK  ( Oryx  gazdla) . 

harnessed  antelopes  (Tragelaphus),  in  which  the  spiral  twisting  of 
the  horns  is  less  marked  than  in  the  other  two ;  these  antelopes 
frequenting  forest  or  jungle,  and  being  most  numerous  in  western 
Africa.  The  kudus  (Strepsiceros]  agree  with  the  last  in  that  the 
females  are  hornless,  but  the  horns  of  the  males  form  a  more 
corkscrew-like  spiral  than  is  generally  the  case  with  the  harnessed 


VII.]  UNGULATES.  247 

antelopes;  while  in  the  elands  (Orias)1  the  horns  are  present  in 
both  sexes  and  form  a  close  spiral. 

Turning  to  the  perissodactyle  section  of  the  order,  we  find 
Ethiopian  Africa  the  home  of  several  species  differing  markedly 
from  any  now  living  in  other  parts  of  the  world,  although,  accord- 
ing to  the  system  here  adopted,  these  do  not  constitute  generic 
groups  by  themselves.  There  are  two,  or  possibly  three  species  of 
Rhinoceros,  both  of  which  are  two-horned,  and  differ  from  those  of 
the  Oriental  region  in  the  absence  of  canine  and  incisor  teeth.  Of 
these  the  common  African  rhinoceros  (R.  bicornis)  is  closely  allied  to 
an  extinct  species  from  the  Pikermi  beds  of  Attica;  while  Burchell's 
rhinoceros  (R.  simus),  which  is  now  nearly  exterminated,  has  its 
nearest  allies  in  the  extinct  R.  platyrhinus  of  the  Siwalik  Hills,  and 
the  woolly  rhinoceros  (R.  antiquitatis)  of  the  Plistocene  of  Europe 
and  northern  Asia.  In  addition  to  being  the  habitat  of  the  parent 
form  of  the  domestic  ass,  which  is  confined  to  Somaliland  and  the 
adjacent  regions,  Ethiopian  Africa  is  also  characterised  by  contain- 
ing all  the  striped  horses,  or  zebras,  separated  by  some  authorities 
as  a  distinct  genus,  under  the  name  of  Hippotigris.  The  best 
known  representatives  of  this  group  are  the  true  or  mountain 
zebra  (Equus  zebra],  Burchell's  zebra  (E.  burcheili\  Grevy's  zebra 
(E.  grevyi]  of  the  Galla  country,  and  the  quagga  (E.  quagga).  The 
latter,  although  formerly  abundant  in  the  southern  extremity  of  the 
continent,  is  now  fast  verging  on  extinction,  and  serves  to  connect 
the  more  typical  members  of  the  group  with  the  true  asses.  In 
spite  of  the  difference  in  their  coloration,  the  zebras  are  indeed 
indistinguishable  in  their  osteology  and  dentition  from  the  latter, 
and  it  is  quite  possible  that  they  are  represented  in  a  fossil  state  in 
the  later  Tertiaries  of  Europe,  while  they  are  not  improbably  the 
direct  descendants  of  the  ancestral  genus  Hipparion.  The  absence 
of  tapirs  from  the  Ethiopian  region  is  as  remarkable  as  the  want 
of  deer ;  but  it  is  noteworthy  that  the  former  are  unknown  among 
both  the  Pikermi  and  Siwalik  faunas. 

Although  the  African  elephant  (Elephas  africanus)  is  markedly 
distinct  from  the  Oriental  species,  yet  the  two  are  so  closely  con- 
nected by  intermediate  extinct  forms  that  it  is  impossible  to  regard 

1  The  name  is  usually  spelt  Oreas,  but  as  it  is  derived  from  <5/>etds,  the 
proper  orthography  is  Orias. 


248 


THE   ETHIOPIAN    REGION. 


[CHAP. 


them  as  the  representatives  of  separate  genera.  The  existing 
species  is  now  confined  to  the  Ethiopian  region,  but  since  its 
fossilised  remains  occur  in  deposits  of  Plistocene  age  in  Algeria, 
Spain,  and  Sicily,  it  is  evident  that,  like  the  spotted  hyaena  and 
lion,  it  formerly  enjoyed  a  much  more  extensive  range. 

With  the  exception  that  a  single  species  is  found  in  Syria,  the 
small  rodent-like  ungulates,  known  as  hyraces,  which  constitute 
not  only  a  family  (Procavtidce)  but  likewise  a  sub-order  (Hyracoidea) 
by  themselves,  are  especially  characteristic  of  the  Ethiopian  region, 
where  they  are  represented  by  a  large  number  of  species,  more 


FIG.  59.     CAPE  HYRAX  (Procavia  capensis]. 

particularly  in  the  southern  portion  of  the  continent.  Although 
these  animals  closely  resemble  the  rhinoceroses  in  the  structure  of 
their  molar  teeth,  they  differ  markedly  from  all  the  perissodactyles 
in  having  the  carpus  constructed  on  the  linear  type1,  and 
from  all  other  living  forms  of  the  order  in  that  their  single  pair 
of  upper  incisor  teeth  grow  continuously  throughout  life,  as  in 
the  rodents.  They  were  formerly  divided  into  at  least  two 
generic  groups,  but  both  the  terrestrial  and  arboreal  forms  are  now 
included  in  the  single  genus  Procavia.  Nothing  definite  is 

1  See  figure  12  on  p.  78. 


VII.]  UNGULATES.  249 

known  as  to  the  past  history  of  the  group1;  but  it  has  been 
suggested  (p.  85)  that  they  may  be  allied  to  certain  extinct  South 
American  ungulates. 

The  list  of  peculiar  Ethiopian  mammals  is  brought  to  a  close 
by  the  aard-varks  (Orycteropodida),  which  although  generally  in- 
cluded in  the  Edentata  have  nothing  to  do  with  the  typical  South 
American  representatives  of  that  order,  and  are  here,  together  with 
the  pangolins,  regarded  as  forming  an  ordinal  group — Effodientia — 
by  themselves.  Only  a  single  genus  (Orycteropus]  now  exists,  of 
which  there  are  two  living  Ethiopian  species,  and  there  are 
extinct  species  in  the  Pliocene  of  Persia  and  Samos.  A  skull 
from  the  Plistocene  of  Madagascar  has  been  described  as  Plesi- 
orycteropus,  and  another  genus  occurs  in  the  French  Oligocene. 
Not  improbably  some  members  of  the  family  entered  Africa  and 
Madagascar  with  the  ancestral  lemuroids  and  civets,  but  the  dis- 
covery of  the  Pliocene  forms  renders  it  probable  that  the  existing 
genus  is  a  later  immigrant. 

Finally,  it  may  be  mentioned  that  among  the  more  widely- 
spread  genera  a  few  species  of  mammals  are  either  now  common 
to  the  Ethiopian  region  and  India,  or  were  so  during  the  Plistocene 
age.  In  the  Felida  the  lion  (Felts  leo),  leopard  (F.  pardus),  jungle- 
cat  (F.  chaus],  caracal  (F.  caracal),  and  hunting-leopard  (Cynce- 
lurus  jubatus]  still  range  over  the  two  areas ;  fossilised  remains  of 
the  first  three  of  these  also  occurring  in  the  European  Plistocene 
deposits.  On  the  other  hand,  the  spotted  hyaena  (Hycena  crocutd), 
which  lived  in  Southern  India  (as  well  as  in  Europe)  during  the 
Plistocene  era,  is  now  restricted  to  Ethiopian  Africa;  and  the 
same  is  the  case  with  the  giant  pangolin  (Mains  gigantea)  of  West 
Africa,  fossilised  remains  of  which  have  been  discovered,  in 
company  with  those  of  the  spotted  hyaena,  in  a  cavern  in 
Madras. 

The  following  table  shows  the  genera  and  family  of  mammals 
now  more  or  less  exclusively  restricted  to  the  Ethiopian  region  ; 
the  names  of  such  as  are  practically  peculiar  to  this  area  being 
printed  in  italic  type. 

1  I  am  informed  that  a  skull  belonging  to  an  extinct  member  of  this  group 
has  been  discovered  in  the  Pliocene  of  Samos,  but  no  description  has  been 
published. 


250  THE    ETHIOPIAN    REGION.  [CHAP. 

I.     Primates. 

1.  ANTHROPOIDEA. 

SlMIID/E. 

Anthropopithecus.     Equatorial  Africa.     Fossil  in 

Indian  Pliocene. 
Gorilla.     W.  African. 

CERCOPITHECID^E. 
Co  lob  us. 

Cercopithecus.     Largely  W.  African. 
Cercocebus.     W.  African. 
Theropithecus.     N.  E.  African. 
Papio.     Fossil  in  Indian  Plistocene  and  Pliocene. 

2.  LEMUROIDEA. 

LEMURID^E. 

Galago.     Equatorial  and  E.  African. 
Perodicticiis.     W.  African. 

II     Insectivora. 

MACROSCELIDID&.     Fossil  in  European  Oligocene. 
Macroscelides  (including  Petrodromus}.     One  N. 

African  species. 
Rhynchocyon.     E.  African. 

ERINACEID^:. 

Proechinus.     W.  African. 


Myosorex. 

POTAMOGALID^:.     Elsewhere  only  in  Madagascar. 

Potamogale.     W.  African. 
CHR  YSOCHLORID&. 

Chrysochloris   (including    Chalcochloris).     S.   and 
E.  African. 

III.     Carnivora. 
FELID^E. 

Cynaelurus.     Elsewhere  only  in  India,  the  same  species 
being  common  to  the  two  regions. 


VII.]  LIST   OF   THE   FAUNA.  251 

III.     Carnivora  (cont.}. 

VlVERRID^. 

Genetta.     One  species  in  South  Holarctic  region. 
Poiana.     W.  African. 
Nandinia.     W.  and  E.  African. 
Helogale.     W.  and  E.  African. 
Bdeogale.     W.  and  E.  African. 
Cynictis.     S.  African. 
Rhynchogale.     E.  African. 
.  Crossarchus. 
Suricata.     S.  Africa. 


Proteles.     S.  and  E.  African. 


Lycaon.     S.    and    E.    African.      Fossil   in    European 

Plistocene.  f 

Otocyon.     S.  African. 

MUSTELID^E. 

Mellivora.     One  African  and  one  Indian  species  ;  and 

another  from  the  Indian  Pliocene. 
Ictonyx.     Ranges    into    Egypt,    and    perhaps    Asia 

Minor. 

Pcecilogale.     S.  African. 
Galeriscus.     E.  African. 

IV.     Rodentia. 

ANOMALURIDM. 

Anomalurus.     W.  and  E.  African. 
Idinrus.     W.  African. 

SCIURID/E. 

Nannosciurus.     Elsewhere  in  Malaysia. 
Xerus.     Fossil  in  European  Miocene. 

MYOXID/E. 

Graphiurus. 

Claviglis.     W.  African.     The  right  of  the  one  species 
to  generic  distinction  is  doubtful. 


252  THE    ETHIOPIAN    REGION.  [CHAP. 

IV.     Rodentia  (cont.\ 


Pachyuromys. 
Mystromys.     S.  African. 
Otomys.     S.  E.  and  W.  African. 
Dasymys.     S.  African. 
Malacomys.     W.  African. 
Dendromys.     \ 

Limacomys.          These  represent  a  peculiar  Ethiopian 
Steatomys.  sub-family  —  the  Dendromyina. 

Lophuromys.    j 

Trilophomys.     N.  E.  African,  and  (?)  S.  Arabian. 
Deomys.     W.  African.     Alone  represents  a  sub-family. 
Cricetomys. 

Saccostomns.     W.  African1. 
SPALACID.E. 

Bathyergus.         \    Constitute    a  sub-family  —  the  Ba- 
Georychus.  thyergina  ;  the  last  of  the  four 

Myoscalops.  being  confined  to  Somaliland, 

Heterocephalus.   )          while  the  first  is  S.  African. 


Pedetes.     The  sole  representative  of  a  sub-family. 
OCTODONTID^:.     Elsewhere  at  the   present  time    only  in 

the  Neogaeic  realm  and  N.  Africa. 
Pectinator.     N.  E.  African. 
Petromys.     S.  African. 
Triaulacodus.     W.,  S.  and  E.  African. 

V.     Ungulata. 

n    Eur°Pean   and    Asiatic 


ARTIODACTYLA 

AK1.     UACiYLA.  Plistocene    and   Pliocene,  and 

HIPPOPOTAMI  DAI.       4  .     ,,    ,  -r.  , 

also  in  Madagascar.     Formerly 


Hippopotamus.  in  lower  Egypt. 


Sus  ;  the  Potamochczrine  group,  frequently  regarded  as 
a  distinct  genus,  is  peculiar  to  the  Ethiopian  and 
Malagasy  regions. 

Phacochcerus. 
1  If  Arvicanthis  be  accepted  as  a  genus  it  should  come  here. 


VII.]  LIST   OF   THE   FAUNA.  253 

V.     Ungulata  (cont.\ 
TRAGULID^E. 

Dor  cat  her  ium.  W.  African  ;  fossil  in  European  Miocene, 

and  also  in  the  Pliocene  of  India. 

GIRAFFID/E.  (  Fossil  in  lower   Pliocene   of  Europe   and 
GiraffaA         Asia. 


Bubalis  (including  Damaliscus}.     Ranges  into  Syria 

and  Tunis  ;  fossil  in  Indian  Pliocene. 
Connochcztes. 
Cephalophus. 

Madoqua.     N.  E.  and  E.  African. 
Nanotragus.     W.  African. 
Nesotragus. 
Rhapiceros. 
Oribia.     S.  African. 
Oreotragus.     S.  and  E.  African. 
Dorcatragus.     Somaliland. 
Pelea.     S.  African. 
Cobus.     Fossil  in  Indian  Pliocene. 
Cervicapra.     S.  W.  and  E.  African. 
sEpyceros.     S.  and  W.  African. 
Ammodorcas.     Somaliland. 
Lithocranius.     E.  African. 
Hippotragus.     Fossil  in  Indian  Pliocene. 
Oryx.     Ranges  into  Syria. 
Tragelaphus.     Perhaps  fossil  in  European  Pliocene. 

Strepsiceros.  ) 

-,   .  f  Fossil  in  Indian  Pliocene. 

Onas.  I 

PERISSODACTYLA. 
RHINOCEROTID^E. 

Rhinoceros  ;  the  species  without  front  teeth  are  now 
peculiar  to  the  Ethiopian  region,  although  allied 
forms    occurred   in    the    European    and    Asiatic 
Pliocene  and  Plistocene. 
EQUID/E. 

Equus  ;  all  the  striped  species  confined  to  this  region. 


254  THE    ETHIOPIAN    REGION.  [CHAP. 

V.     Ungulata  (cont.). 
3.     HYRACOIDEA\ 

PROCAVIID/E.      I  Range  into  Syria. 
Procavia. 

VL     Effbdientia.     Ethiopian  and  Oriental. 

ORYCTEROPODIDJS.     Fossil  in  French  Oligocene. 

Orycteropus.     Fossil  in  Pliocene  of  Samos  and  Persia. 

Among  groups  other  than  mammals,  attention  may  be  directed 
to  the  remarkable  difference  between  the  birds  of 
Ethiopia  and  those  of  Madagascar.  On  this  point 
Dr  Blanford1  writes  that  "the  most  characteristic  African  families, 
such  as  plantain-eaters  (Musophagidce),  colies  (Collidce),  and  wood- 
hoopoes  (Irrisoridce),  barbets,  hornbills,  secretary-birds  (Serpen- 
tarius],  and  a  number  of  genera,  such  as  Lamprotornis  (glossy 
starlings),  Buphaga  (ox-peckers),  Laniarius,  and  Telephonus,  that 
are  the  common  and  familiar  birds  of  every  part  of  Africa  south  of 
the  Sahara,  are  entirely  wanting  in  the  Mascarene  Islands,  in- 
cluding the  Seychelles,  Mauritius,  etc.,  while  no  fewer  than  four 
peculiar  families  and  a  number  of  genera  confined  to  the  archi- 
pelago replace  them.  Amongst  the  Mascarene  birds,  too,  are 
found  several  representatives  of  Oriental  genera,  or  genera  closely 
allied  to  Oriental  types,  and  without  any  near  Ethiopian  relations. 
Foremost  among  these  are  certain  bulbuls,  forming  the  genera 
Ixocincla  and  Tylas,  the  former  composed  of  species  which  have 
been  usually  referred  to  the  typically  Oriental  genus  Hypsipetes, 
and  the  latter  nearly  affined.  In  fact,  as  was  shown  by  Geoffrey 
St  Hilaire,  and  as  Hartlaub  has  since  pointed  out,  there  is  in  the 
Mascarene  avifauna  a  more  marked  connexion  with  Indian  than 
with  Ethiopian  types.  In  the  Seychelles,  especially,  out  of  the 
seven  Passerine  genera  represented  by  peculiar  species,  three, 
Nectarinia,  Zosterops,  and  Tchitrea,  are  Indian  and  African,  one, 
Foudia,  is  Ethiopian,  but  not  Indian,  and  two,  Copsychns  and 
Hypsipetes,  or  Ixocincla,  are  Indian  but  not  African." 

All  this  is  confirmatory,  not  only  of  the  right  of  Madagascar 
and  the  Mascarenes  to  form  a  region  by  themselves,  but  likewise 

1  Appendix,  No.   8,  p.  89.     In  this  quotation  the  English  or  Latin  names 
have  in  some  cases  been  added  to  the  original. 


VII.]  PAST   HISTORY.  255 

of -the  distinction  between  the  Ethiopian  and  Oriental  regions, 
which  some  have  proposed  to  unite.  It  is,  however,  somewhat 
remarkable  that  secretary-birds  (Serpentarius)  are  unknown  in 
Madagascar,  seeing  that  they  are  represented  in  the  upper  Oligo- 
cene  of  France,  and  may  therefore  be  presumed  to  have  entered 
Ethiopia  with  the  ancestral  lemuroids  and  civets.  Finally,  the 
ostriches  (Struthio),  which  are  now  mainly  confined  to  Africa  and 
Syria,  are  evidently  recent  immigrants  into  the  region,  the  genus 
being  represented  in  a  fossil  state  in  the  Indian  Pliocene. 

With  the  exception  of  the  occurrence  of  remains  of  certain 
existing  species,  such  as  Rhinoceros  simus,  Phaco- 
charus,  etc.,  in  the  superficial  deposits  of  southern 
Africa,  nothing  is  known  of  the  mammalian  Tertiary 
palaeontology  of  the  Ethiopian  region.  Fortunately,  however,  the 
clue  given  by  the  existing  fauna  of  Madagascar  and  the  Tertiary 
faunas  of  Europe  and  southern  Asia  enables  a  considerable  portion 
of  the  past  history  of  the  population  of  the  region  to  be  given  with 
a  fair  degree  of  completeness.  And  here,  with  one  important 
exception,  Dr  Wallace's  explanation,  as  given  in  the  Geographical 
Distribution  of  Animals1  >  may  be  accepted  almost  in  its 
entirety; — the  one  exception  being,  as  mentioned  in  an  earlier 
chapter,  that  the  Sahara  was  never  a  sea  during  Tertiary  times ; 
although  it  appears  always  to  have  formed  a  barrier  between 
northern  Africa  and  Ethiopia.  As  already  mentioned,  the  an- 
cestral types  of  the  existing  mammalian  fauna  of  Madagascar  ob- 
tained an  entrance  into  Ethiopia  some  time  during  the  Oligocene 
period,  and  soon  after  ranged  over  the  whole  of  what  are  now  the 
Ethiopian  and  Malagasy  regions,  which  were  then  united  and 
possessed  a  common  fauna.  During  the  Pliocene  age,  when 
Madagascar  had  become  isolated,  came  the  great  irruption  into 
Ethiopia,  of  the  higher  and  larger  mammals,  such  as  apes,  monkeys, 
ungulates,  etc.,  which  were  then  flourishing  all  along  southern 
Europe  and  Asia.  Finding  the  country  unoccupied  and  eminently 
suited  to  their  existence,  these  rapidly  attained  a  development  now 
unequalled  in  any  other  part  of  the  world  ;  many  new  genera  being 
apparently  evolved  within  the  Ethiopian  area,  although  a  large 

1  Vol.  i.  p.  285 — 292. 


256  THE    ETHIOPIAN    REGION.  [CHAP. 

number  were  already  in  existence  at  the  time  of  the  southern 
migration.  Several  of  these  existing  genera  are  met  with  in  the 
Pikermi  deposits  of  Greece,  but  more  were  confined,  at  this  epoch, 
to  the  Pliocene  of  Persia,  Samos,  and  India ;  and  it  may  there- 
fore be  assumed  that  the  great  migration  was  by  way  of  Syria  or 
Arabia.  Dr  Wallace  has  indeed  expressed  the  opinion  that  a 
certain  number  of  types — among  them  the  elephants  and  rhinoce- 
roses— obtained  an  entrance  to  the  westward  of  Tunis  ;  but  there 
are  no  true  elephants  in  the  Pikermi  deposits,  and  apparently 
none  in  those  of  Persia,  whereas  their  remains  abound  in  the 
Siwalik  Hills.  As  to  the  rhinoceroses,  although  the  Pikermi 
species  is  closely  allied  to  the  African  Rhinoceros  bicornis,  the 
Siwalik  R.  platyrhinus  is  equally  close  to  R.  simus ;  and  in  the 
Siwaliks  we  meet  with  chimpanzees  (Anthropopithecus),  baboons 
(Papio),  ratels  (Mellivora),  hippopotami,  water-chevrotains  (Dorca- 
therium\  and  several  genera  of  Ethiopian  antelopes,  all  of  which 
are  totally  unknown  in  the  Pikermi  beds.  Ostriches,  too,  are 
first  known  in  the  Siwaliks ;  while  aard-varks  occur  in  the  Persian 
and  Samos  beds.  All  the  evidence  accordingly  points  to  the 
great  immigration  having  taken  place  along  the  eastern  side  of  the 
continent ;  and  the  existence  of  certain  species  of  mammals  which 
are  either  still  common  to  India  and  Africa,  or  which  were  so 
during  the  Plistocene  epoch,  lends  support  to  this  view.  Further 
testimony  in  this  direction  is  aiforded  by  the  occurrence  of  closely- 
allied  generic  types  in  the  Ethiopian  and  Oriental  regions. 
Among  the  lemuroids,  for  instance,  the  Oriental  lorises  (Nycti- 
cebus  and  Loris)  are  replaced  in  Western  Africa  by  the  potto  and 
awantibo  (Perodicticus) ;  while  in  the  Viverrida  the  true  linsangs 
(Linsanga)  of  the  eastern  half  of  the  Oriental  region  are  repre- 
sented in  Fernando  Po  by  the  allied  Poiana,  and  the  Oriental 
palm-civets  (Paradoxurus}  have  very  close  allies  in  the  two  species 
of  the  Ethiopian  genus  Nandinia.  A  less  marked  instance  is 
afforded  by  the  occurrence  of  the  water-chevrotain  (Dorcatheriuni} 
in  West  Africa  and  of  the  true  chevrotains  in  southern  India  and 
the  eastern  half  of  the  Oriental  region. 

And  here  it  may  be  remarked  that  especial  stress  has  been  laid 
upon  the  much  greater  resemblance  that  exists  between  the  fauna 
of  the  eastern,  or  Malayan,  division  of  the  Oriental  region  and 


VII.]  ETHIOPIAN    AND   MALAYAN    FAUNAS.  257 

Western  Africa,  than  between  that  of  peninsular  India  and 
Eastern  and  South  Africa  ;  large  man-like  apes  and  linsangs  being 
confined  in  the  Oriental  region  to  its  eastern  half,  while  palm- 
civets,  lorises,  and  chevrotains  are  more  abundant  there  than  in 
other  parts  of  the  same  region.  This,  however,  appears  to  be 
mainly  or  entirely  due  to  similarity  of  climatic  conditions,  and 
not  to  original  distributional  distinctions.  And,  it  may  be  re- 
marked, increased  acquaintance  with  the  fauna  of  Ethiopia  tends 
to  show  that  types  formerly  thought  to  be  confined  to  the  western 
half  of  Africa  really  extend  far  to  the  eastward ;  chimpanzees,  for 
instance,  being  now  known  to  range  as  far  east  as  Ugogo,  while 
the  genus  Nandinia,  which  was  originally  known  solely  by  a  West 
African  species,  is  now  proved  to  have  an  eastern  representative  in 
Nyasaland. 

The  similarity  between  the  fauna  of  the  Malayan  sub-region 
and  that  of  Western  Africa  naturally  leads  on  to  the  consideration 
of  the  former  land-connection  between  India  and  Africa.  Writing 
on  this  subject,  Dr  Wallace1  observes  that  "we  may  now  perhaps 
see  the  reason  of  the  singular  absence  from  tropical  Africa  of  deer 
and  bears ;  for  these  are  both  groups  which  live  in  fertile  and  well- 
wooded  countries,  whereas  the  line  of  immigration  from  Europe  to 
Africa  was  probably  always,  as  now,  to  a  great  extent  a  dry  and 
desert  tract,  suited  to  antelopes  and  large  felines,  but  almost 
impassable  to  deer  and  bears.  We  find,  too,  that  whereas  remains 
of  antelopes  and  giraffes  abound  in  the  Miocene2  deposits  of 
Greece,  there  were  no  deer  (which  are  perhaps  a  somewhat  later 
development),  neither  were  there  any  bears,  but  numerous  forms 
of  Felidce,  Viverrida,  Mustdida,  and  ancestral  forms  of  Hyczna, 
exactly  suited  to  be  the  progenitors  of  the  most  prevalent  types  of 
modern  African  zoology." 

As  mentioned  in  an  earlier  chapter,  since  this  passage  was 
written  the  discovery  of  the  remains  of  a  species  of  chimpanzee 
in  the  Indian  Siwaliks  has  shown  quite  clearly  that  the  line 
of  communication  between  India  and  Africa  must  have  included 
a  wooded  tract  comparable  to  the  existing  equatorial  African 
forest  region ;  and  this  would  be  true  even  if  the  migration 

1  Op.  dt.  p.  291. 

2  The  Pikermi  beds  were  formerly  universally  held  to  be  of  Miocene  age. 

L.  I 


258  THE   ETHIOPIAN    REGION.  [CHAP. 

had  taken  place  from  Africa  to  India,  which  was  not  the 
case.  Evidence  of  such  a  tract  is,  I  believe,  afforded  by  the 
occurrence  of  fossilised  tree-stems  in  many  districts  which  are  now 
desert.  And  along  this  tract  there  can  be  little  doubt  that  the 
ancestors  of  the  mammalian  types  now  common  to  the  West 
African  and  Malayan  sub-regions  originally  wandered  from  their 
common  Indian  home.  Subsequently  the  whole  of  the  countries 
lying  between  eastern  Africa  and  India  have  become  deforested, 
while  in  Africa  itself  the  forest-area  has  shrunk  away  from  the 
eastern  side  of  the  continent. 

This  leaves  the  question  of  the  absence  of  bears  and  deer  from 
Africa  without  any  adequate  explanation.  Bears  are,  however,  in 
the  main,  mountain  animals,  some  of  which,  like  the  isabelline 
bear  of  the  Himalaya,  inhabit  districts  where  there  is  but  little 
forest ;  and  it  is  noteworthy  that,  with  the  exception  of  the  sloth- 
bear,  which  forms  a  genus  apart  (Melursus)^  there  are  no  bears  in 
India  proper,  although  a  fossil  species  allied  to  the  sloth-bear 
occurs  in  the  Siwaliks.  This  being  so,  when  we  take  into  account 
the  absence  of  ursine  remains  from  the  Pikermi  and  Persian  beds, 
there  is  nothing  very  wonderful  in  the  fact  that  none  of  these 
animals  entered  Ethiopia  during  the  great  Pliocene  migration. 
The  absence  of  all  Cervidce  is  more  difficult  to  explain,  seeing 
that  deer  of  an  Oriental  type  are  abundant  in  the  Siwaliks,  while 
they  are  also  sparingly  represented  in  the  Pikerrni  beds.  Typical 
deer  of  the  red-deer  group  are,  however,  totally  wanting  in  the 
Siwaliks,  as  they  are  in  the  Oriental  region  at  the  present  day ; 
and  we  are,  therefore,  perfectly  able  to  account  for  their  absence 
from  the  Ethiopian  region,  although  they  occur  in  Africa  north  of 
the  Sahara.  With  regard  to  the  absence  of  Oriental  types  of  deer 
in  Ethiopia,  it  can  only  be  said  that  it  is  as  difficult  to  see  any 
reason  why  these  should  have  continued  to  flourish  since  the 
Pliocene  in  the  Oriental  region  without  ever  having  entered  Africa, 
as  it  is  to  explain  why  giraffes,  hippopotami,  and  ostriches  should 
have  disappeared  from  the  former  area  to  survive  in  the  latter. 

Another  difficulty  is  presented  by  the  case  of  the  pigs,  bat  here 
it  may  be  suggested  that  the  absence  of  the  typical  group  of  the 
genus  Sus  from  the  whole  of  Ethiopia,  with  the  exception  of 
Sennaar,  may  perhaps  be  accounted  for  by  all  the  other  species  of 


VII.]  LEADING   CHARACTERISTICS.  259 

that  genus  which  originally  entered  the  country  having  been 
developed  into  the  more  specialised  Phacochxrus.  Attention  has 
already  been  called  to  the  fact  that  the  molars  of  some  of  the 
Indian  Tertiary  species  of  Sus  show  a  distinct  approximation  to 
those  of  Phacochairus,  and  further  evolution  might  easily  lead  to 
the  development  of  the  latter.  Since  this  genus  is  unknown  in  a 
fossil  state  from  other  regions,  is  it  improbable  that  it  has  originated 
from  Sus  within  the  limits  of  its  present  habitat  ? 

Summarising  the  results  of  the  foregoing  survey  of  the  mam- 
malian fauna  of  Ethiopia,  it  would  appear  that  the  Sahara  has  for 
a  very  long  period  formed  a  barrier  between  Ethiopian  Africa  and 
the  northern  part  of  the  continent.  When  first  populated  by 
Tertiary  mammals,  Ethiopia  and  Madagascar  were  in  union,  and 
formed  but  a  single  zoological  province,  which  would  seem  to 
have  been  to  a  great  extent  isolated  from  the  rest  of  the  Old 
World  during  the  Miocene  epoch.  Had  such  conditions  persisted 
|his  province  would  have  been  entitled  to  form  a  primary 
zoological  realm  by  itself.  During  the  Pliocene  epoch,  however, 
Madagascar  became  separated,  while  a  more  complete  union  of 
the  continent  with  Asia  by  way  of  Syria  or  Arabia  permitted  the 
influx  of  larger  and  higher  mammals  from  the  eastward.  Hence 
there  is  a  most  intimate  relationship  between  the  Pliocene  fauna 
of  India  and  the  one  now  inhabiting  Ethiopia ;  but  the  distinc- 
tion between  the  two  areas  at  the  present  day  is  fully  sufficient 
to  justify  the  separation  of  the  Ethiopian  from  the  Oriental  region. 
Of  all  the  zoological  regions  of  the  world,  the  Ethiopian  may  be 
regarded  as  the  one  which  has  been  most  recently  evolved ;  and 
it  may  be  shortly  characterised  by  the  sole  possession  of  the 
gorilla  and  chimpanzees ;  the  absence  of  bears  and  deer ;  and  the 
presence  of  the  African  elephant,  hyraces,  rhinoceroses  devoid  of 
front  teeth,  zebras,  hippopotami,  wart-hogs,  giraffes,  numerous 
genera  of  antelopes,  and  aard-varks,  as  well  as  by  the  great  de- 
velopment of  its  large  ungulates  in  general.  It  shares  with  the 
Oriental  region  the  distinction  of  being  the  sole  habitat  at  the 
present  day  of  man-like  apes,  true  civets  ( Viverra),  linsangs,  palm- 
civets1,  ratels  (Mellivora),  elephants,  rhinoceroses,  and  pangolins 

1  Also  in  the  Austro-Malayan  region. 

17—2 


260  THE   ETHIOPIAN    REGION.  [CHAP. 

(Mam's),  together  with  the  rodent  genera  Nannosciurus,  Golunda 
and  Atherura.  The  probable  relationship  between  Ethiopia  and 
Neogaea  has  been  sufficiently  discussed  in  the  third  chapter. 

Although,  from  the  mammalian  point  of  view,  Ethiopia  is  a 
very  modern  region,  as  a  continent  it  is  one  of  the  oldest,  the 
greater  portion  of  its  area  having  been  land  since  the  Palaeozoic 
epoch.  As  has  been  shown  in  an  earlier  chapter,  in  Palaeozoic 
times  southern  Africa  formed  a  portion  of  the  great  southern  or 
equatorial  continent  distinguished  from  more  northern  lands  by 
the  peculiar  characteristics  of  its  flora;  and  it  is  probable  that 
it  remained  connected  with  India  until  late  in  the  Secondary 
epoch1;  as  is  proved  by  the  identity  of  the  flora  and  reptiles  of  the 
two  areas.  Early,  however,  in  this  epoch  there  must  also  have 
been  free  communication  with  Europe,  as  shewn  by  the  close 
alliance  of  the  anomodont  reptiles  of  the  two  continents,  and 
likewise  by  the  occurrence  in  both  of  the  mammalian  genus 
Tritylodon.  In  the  Cretaceous,  so  far  as  vertebrates  are  conf 
cerned,  our  knowledge  of  the  Ethiopian  region  is  practically  a 
blank. 

In  conformity  with  the  plan  adopted  in  other  cases,  the  sub- 
regions  into  which  Ethiopian  Africa  may  be  divided 

Sub-regions.  .  . 

will  be  treated  very  briefly. 

Writing  of  the  desert-tracts  of  the  Saharan  sub-region,  where 
the  necessary  conditions  for  existence  are  largely  wanting,  Dr 
Heilprin  observes  that  "  there  is  a  marked  impoverishment  of  the 
fauna.  The  more  formidable  carnivores,  such  as  the  lion  and  the 
leopard,  are  absent  from  most  districts,  leaving  their  places  to  be 
filled  by  some  minor  cats,  the  hyaena,  jackal,  fox,  and  fennec. 
The  hoofed  animals  are  represented  (in  some  parts)  by  the  buffalo, 
and  a  limited  number  of  antelopes  (Gazella,  Oryx,  and  Addax). 
Among  rodents  the  families  of  rats  (Muridce)  and  jumping-mice 
(Dipodidcd)  are  fairly  represented,  in  addition  to  which  we  have 
the  porcupine  and  hare  (Lepus  mediterraneus}.  The  ostrich  is 
sufficiently  abundant  throughout  most  of  the  sub-region." 

In  marked  contrast  to  the  poverty  of  the  Saharan  districts, 
is  the  remarkable  richness  of  the  great  equatorial  forest-tract, 

1   Vide  siipra,  p.  224. 


VII.]  SUB-REGIONS.  26l 

which  is  the  exclusive  home  of  all  the  man-like  apes  and  of  the 
lemurs  of  the  genus  Perodicticus.  To  this  sub-region  also  belong 
PotamogaUi  the  African  linsang  (Poiana),  as  well  as  several  other 
genera  of  Viverridcz,  such  as  Nandinia  and  Helogale.  Among  the 
rodents  the  flying-squirrels  of  the  family  Anomalurida  are  very 
characteristic  of  the  forest  tract,  and  the  peculiar  murine  genus 
Deomys,  as  also  Saccostomus,  is  restricted  to  it.  The  water-chevro- 
tain  (Dorcatheriuni)  is  solely  West  African,  as  is  the  small  Liberian 
hippopotamus ;  while  certain  genera  of  antelopes,  such  as  the 
duikers  (Cephalophus),  harnessed  antelopes  (Tragelaphns\  and 
elands  (Orias),  have  larger  and  finer  representatives  here  than 
elsewhere.  The  giant  pangolin  (Manis  gigantea),  the  largest 
member  of  its  genus,  is  likewise  a  West  African  form. 

Although  South  Africa  has  a  certain  number  of  mammalian 
genera,  such  as  Cynictis,  Suricata,  Otocyon,  Pcecilogale,  Bathyergus, 
Pedetes,  Petromys,  and  Pelea,  peculiar  to  it,  others,  such  as  the 
golden  moles  (Chrysochloris)  and  aard-wolf  (Proteles],  have  been 
proved  to  extend  far  up  the  east  coast,  the  latter  occurring  in 
Somaliland.  Hence  it  seems  advisable  to  unite  Dr  Wallace's 
South  African  sub-region  with  that  portion  of  his  East  Central 
sub-region  which  is  not  included  in  the  equatorial  forest-tract.  Of 
this  East  Central  sub-region,  as  it  may  be  collectively  called,  it 
will  suffice  to  say  that  it  is  the  home  of  the  greater  number  of  the 
characteristic  Ethiopian  mammals  exclusive  of  those  restricted  to 
the  forest-tract.  Here  antelopes  attain  their  greatest  numerical 
development,  both  as  regards  genera  and  species;  and  here  is 
also  the  true  home  of  the  lion  and  the  spotted  hyaena ;  while  to 
this  sub-region  are  confined  the  Cape  hunting-dog  (Lycaon)  and 
the  aard-wolf  (Proteles).  The  distribution  of  other  genera  is 
sufficiently  indicated  in  the  table  already  given. 

The  light  which  has  recently  been  thrown  upon  the  mammals 
of  northern  Somaliland  has  shown  that,  as  regards  antelopes  at 
least,  these  are  so  peculiar  that  it  may  be  questioned  whether  this 
tract  is  not  entitled  to  be  separated  as  a  sub-region  by  itself. 
According  to  the  lists  given  by  Dr  Sclater1  and  Capt.  Swayne2, 


1  Natural  Science,  vol.  I.  p.  264  (1892). 

2  Seventeen  Trips  to  Somaliland,  London  (1895). 


262 


THE    ETHIOPIAN    REGION. 


[CHAP. 


there  are   no   less   than    sixteen 
northern  Somaliland,  of  which  the 

1.  Bubalis  swaynei. 

2.  Madoqua  swaynei. 

3.  „         phillipsi. 

4.  ,,         guentheri. 

5.  Oreotragus  saltator. 

6.  Dorcatragus  megalotis. 

7.  Cobus  ellipsiprymnus. 

8.  Gazella  pelzelni. 


species  of  antelopes  found  in 
names  are  as  follows,  viz. : 

9.     Gazella  spekei. 

10.  ,,       soemmerringi. 

11.  Ammodorcas  clarkei. 

12.  Lithocranius  walleri. 

13.  Oryx  beisa. 

14.  Tragelaphus  decula. 

15.  Strepsiceros  kudu. 

1 6.  imbubis. 


Among  these  Nos.  i,  2,  3,  4,  6,  9,  n  are  quite  peculiar  to  this 
district,  while  No.  12,  which,  like  Nos.  6  and  n,  is  the  sole  repre- 
sentative of  its  genus,  is  only  found  elsewhere  along  the  east  coast 
as  far  south  as  the  Tana  river.  Another  generic  form  peculiar  to 
Somaliland  is  Heterocephalus,  including  two  small  naked  rodents 
with  burrowing  habits ;  while  in  the  same  order  the  single  species 
of  Pectinator  is  restricted  to  this  district.  Among  species  may  be 
mentioned  two  musk-shrews  (Crocidura  smithi  and  C.  somalica),  a 
hedgehog  (Erinaceus  sclateri),  as  well  as  a  banded  mungoose 
(Crossarchus  somalicus).  The  Somali  ostrich  seems  likewise  to 
represent  a  species  by  itself.  On  the  other  hand,  in  addition  to 
those  mentioned  in  the  foregoing  list  of  antelopes,  there  are 
several  East  or  South  African  mammals,  such  as  the  aard-wolf, 
which  range  into  Somaliland,  and  further  evidence  is  perhaps 
desirable  before  the  right  of  that  country  to  form  a  separate  sub- 
region  can  be  admitted. 

The  case  is  more  clear  with  regard  to  south-eastern  Arabia, 
whence  Mr  O.  Thomas1  records  the  following  fifteen  species  of 
land-mammals,  viz. :  Xantharpyia  amplexicaitdata,  Taphozous  nudi- 
ventriS)  Rhinopoma  microphyllum,  Erinaceus  niger,  Crocidura 
murina,  Herpestes  albicauda,  Cants  pallipes,  C.  leucopus,  Gerbillus 
dasyurus,  Mus  rattus,  Lepus  omanensis,  Gazella  muscatensis,  Oryx 
beatrix,  Hemitragus  jayakari,  and  Procavia  syriaca.  Of  these  Mr 
Thomas  remarks  that  their  geographical  relationships  "  are,  as 
might  be  expected,  about  equally  with  Africa  and  India,  three  of 

1  Proc.  Zool.  Soc.  1894,  p.  449.     In  one  case  the  generic  title  has  been 
altered,  in  order  to  bring  it  into  harmony  with  the  system  followed  in  this  work. 


VII.]  SUB-REGIONS.  263 

the  species  being  distinctly  African  in  affinities,  three  Indian,  and 
the  remainder  either  peculiar  or  widely-spread,  and  of  no  special 
significance."  The  association  of  a  goat  belonging  to  the  Oriental 
genus  Hemitragus  with  such  an  essentially  Ethiopian  animal  as  a 
hyrax  {Procavia},  shows  that  we  are  here  truly  on  the  border-land 
between  the  two  regions  in  question. 

In  conclusion,  from  whatever  aspect  it  be  regarded,  Ethiopia 
is  one  of  the  most  interesting  of  the  regions  of  Arctogaea;  and  if, 
as  is  suggested  in  an  earlier  chapter,  it  has  been  the  feeder  by 
means  of  which  South  America  received  its  earliest  mammalian 
Tertiary  fauna,  it  is  entitled  to  an  importance  above  all  the  other 
zoological  regions  of  the  realm  to  which  it  pertains. 


CHAPTER  VIII. 

THE   ORIENTAL   REGION. 

Sub-regions — Characteristics  of  the  Mammalian  Fauna — Past  History  of  the 
Region — Malayan  sub-region — Nicobars,  Mentawi,  and  Christmas  Islands 
— Philippine  sub-region. 

FAR  inferior  in  extent  to  the  Ethiopian,  the  Oriental  region  is 
taken  to  include  those  portions  of  the  continent  of  Asia  lying 
south  of  the  Holarctic  region  (with  the  exception  of  southern 
Arabia,  which  is  Ethiopian),  together  with  the  islands  of  Ceylon, 
Formosa,  the  Philippines,  Sumatra,  Java,  Borneo,  and  numerous 
smaller  ones.  In  India  the  northern  limits  of  the  region  are 
formed  by  the  higher  ranges  of  the  Himalaya,  while  "  Wallace's 
line"  constitutes  the  eastern  boundary  denning  it  from  the  Austro- 
Malayan  region  of  the  Notogaeic  realm.  In  a  region  so  diversified 
as  the  Oriental,  it  would  not  be  natural  to  expect  a  homogeneous 
fauna;  and,  as  a  matter  of  fact,  there  are  in  this  respect  great 
diversities  between  the  different  portions  of  the  region,  many  of 
the  peculiar  genera  having  a  very  restricted  distribution.  Never- 
theless, the  positive  and  negative  features  of  the  mammalian  fauna 
of  the  region  as  a  whole  are  sufficient  to  indicate  its  zoological 
unity,  and  also  to  differentiate  it  from  the  Ethiopian  region,  to 
which  it  is  now  most  closely  allied.  In  the  Himalaya  there  is  a 
gradual  transition  towards  the  Holarctic  fauna ;  and  it  is  probable 
that  in  this  portion  of  the  area  the  differentiation  between  the 
Oriental  and  Holarctic  faunas  has  been  largely  due  to  the  eleva- 
tion of  the  Himalaya  itself,  which  has  taken  place  entirely  since 
the  early  part  of  the  Tertiary  period,  and  is  to  a  considerable 
extent  of  Post-tertiary  date.  It  has  already  been  shown  that  the 
older  Pliocene  fauna  of  northern  India  and  Burma  contained  a 


CHAP.  VIII.]  PHYSICAL   FEATURES.  265 

remarkable  admixture  of  mammalian  genera  now  respectively 
confined  to  the  Ethiopian  and  Oriental  regions,  together  with 
some  of  a  Holarctic  facies ;  and  the  completion  of  the  elevation 
of  the  Himalayan  chain  was  probably  an  important  factor  in  the 
dispersal  and  differentiation  of  this  common  fauna.  Holarctic 
types  are  again  met  with  in  force  in  the  open  desert  regions  on 
the  north-western  frontier  of  India.  On  the  other  hand,  the 
fauna  of  the  Philippines  exhibits  an  approximation  to  that  of  the 
Austro-Malayan  region,  and  thus  shows  a  blending  between  the 
Arctogaeic  and  Notogaeic  realms.  In  physical  features  the  Oriental 
region  displays  great  variation,  a  large  portion  of  peninsular  India 
consisting  of  open  dry  grassy  plains,  whereas  the  slopes  of  the 
eastern  Himalaya,  together  with  the  greater  part  of  Assam,  Burma, 
and  the  Malayan  countries  are  clad  with  luxuriant  forests ;  these 
tropical  or  sub-tropical  forest-regions  being  those  where  the  fauna 
attains  its  fullest  and  richest  development. 

The  poorest  part  of  the  region  is,  as  Dr  Wallace1  observes,  the 
great  triangular  plateau  forming  the  Indian  peninsula;  this  area 
differing  remarkably  from  the  Himalaya  in  its  geological  features, 
and  having  been  land  since  an  extremely  ancient  date,  whereas 
the  Himalayan  area  consists  very  largely  of  marine  formations. 
Since  it  is  stated  in  the  passage  cited  that  peninsular  India 
during  the  Tertiary  period  existed  as  an  island  entirely  discon- 
nected from  the  Himalaya  and  Burma,  it  may  be  well  to  quote 
the  later  and  more  authentic  views  of  the  authors  of  the  Manual 
of  the  Geology  of  India2  on  this  subject.  After  reference  to  the 
extent  of  Eocene  rocks  in  northern  India,  it  is  there  stated  that 
"  the  Peninsula  of  India  in  Eocene  times  was  part  of  a  tract 
of  land,  perhaps  of  a  great  continent  united  to  Africa ;  that  there 
was  a  sea  to  the  eastward,  extending  far  to  the  north-east,  in  the 
region  now  occupied  by  the  Assam  hills,  and  another  sea  to  the 
north-west,  covering  great  part,  if  not  the  whole,  of  Persia,  Baluch- 
istan, the  Indus  plain,  and  a  portion  of  the  upper  Ganges  plain. 
An  arm  of  this  sea  extended  from  the  north-west  up  the  Indus 
valley  in  Ladak.  The  Himalaya,  and  perhaps  Tibet,  wholly  or 
in  part,  were  raised  above  the  sea ;  but  formed  in  all  probability 

1  Geographical  Distribution  of  Animals,  vol.  I.  p.  314. 

2  First  edition,  pt.  I.  p.  liii. 


266  THE   ORIENTAL   REGION.  [CHAP. 

land  of  moderate  elevation.  Whether  the  Himalayan  land  was 
united  to  the  Peninsula  is,  of  course,  uncertain — but  very  pro- 
bably it  was ;  for  there  is  no  evidence  of  marine  conditions  having 
existed  in  the  Ganges  plain  to  the  east  of  the  Dehra  Dun ;  and  if 
the  ferruginous  bands  of  the  Subathu  group  be  laterite,  as  they 
appear  to  be,  the  trappean  detritus  composing  them  must  have 
been  derived,  in  all  probability,  from  the  peninsular  area;  and  the 
latter  must  consequently  have  extended  northward  to  the  base  of 
the  Himalayas,  in  the  neighbourhood  of  Umballa....  In  Miocene 
times,  although  marine  conditions  prevailed  throughout  western 
Sind,  the  area  of  the  sea  was  very  much  smaller  than  in  the  Eocene 
period ;  for  all  the  marine  beds  of  the  Punjab  and  Sub-Himalayas 
are  destitute  of  marine  fossils,  and  are  probably  fluviatile  de- 
posits." 

From  this  it  would  appear  probable  that  during  the  whole  of 
later  Tertiary  times,  at  least,  there  has  been  no  isolation  of 
peninsular  India  from  the  eastern  Himalaya  and  the  Burmese 
countries ;  and  consequently  that  the  differences  between  the 
faunas  of  these  areas  are  mainly  or  solely  due  to  their  differences 
of  physical  features  and  climate. 

By  Dr  Wallace  the  Oriental  region  is  divided  into  four  sub- 
reerions  ;  namely  (i)  the  Indian,  comprising  the  whole 

Sub-regions.  J  ^    ' 

of  upper  India,  (2)  the  Ceylonese,  including  southern 
India  and  Ceylon,  (3)  the  Indo-Chinese,  embracing  Assam,  Burma, 
and  such  portion  of  China  as  lies  within  the  limits  of  the  region, 
and  (4)  the  Indo-Malayan,  which  includes  not  only  the  Malayan 
archipelago  and  islands,  but  likewise  the  Philippines. 

So  far  as  India  and  its  dependencies  are  concerned,  the  follow- 
ing amended  scheme  has  been  proposed  by  Dr  Blanford l,  viz. : 
i.     Himalayan.     The  southern  slopes  of  the  Himalaya,  from  the 

base  to  about  the  limit  of  trees. 

ii.  Indian.  India  from  the  base  of  the  Himalaya  to  Cape 
Comorin,  with  the  exception  of  the  Malabar  coast,  but  with 
the  addition  of  northern  Ceylon. 

iii.  Malabar  or  Ceylonese.  The  Malabar  coast  and  the  neigh- 
bouring hills  as  far  north  as  the  Tapti  river,  together  with 
southern  Ceylon. 

1  Fatina  of  British  India — Mammalia,  p.  v. 


VIII.]  SUB-REGIONS.  267 

iv.     Burmese.     All  Burma  except  south  Tenasserim,  and  with  the 

addition  of  Assam  and  the  intervening  countries, 
v.     Malayan.     South  Tenasserim,  the  Malay  Peninsula,  and  the 

Malayan  Islands  as  far  as  Wallace's  line. 
Whether  the  Philippine   Islands   should   be  included  in  this 

sub-region,   or  should  form    one    by  themselves,  may  be 

doubtful, 
vi.     Indo-Chinese.     Although  not  free  from   objection,  this  term 

may  be  employed   for    the   sub-region   indicated  by  that 

portion  of  China  coming  within  the  limits  of  the  Oriental 

region. 

Regarding  these  sub-regions  in  general,  Dr  Blanford  observes 
that  some  "  may  require  further  subdivision.  Thus  the  fauna  of 
the  North-west  Provinces  and  Punjab  differs  considerably  from 
that  of  southern  India,  and  both  areas  exhibit  zoological  dis- 
tinctions from  the  forest-clad  tracts  of  south-western  Bengal. 
There  is  also  much  difference  between  the  animals  of  Pegu  and 
Arakan,  on  the  one  hand,  and  those  of  the  drier  regions  of  upper 
Burma  on  the  other ;  and  even  greater  distinctions  may  be  traced 
between  those  found  in  the  sub-tropical  and  those  inhabiting  the 
temperate  regions  of  the  Himalaya.  On  the  other  hand,  the  sub- 
tropical Himalayas  were  united  with  the  Burmese  sub-region  by 
Wallace,  and  the  two  are,  perhaps,  zoologically  more  allied  to 
each  other  than  to  any  other  sub-region." 

Recent  discoveries  clearly  indicate  that  the  Philippine  Islands, 
exclusive  of  Palawan  and  the  Calamianes,  should  form  a  sub-region 
by  themselves. 

Taking  the  Oriental  region  as  a  whole,  it  may  be  stated  that 
the  number  of  peculiar  generic  types  of  mammals  is 
less  than  in  the  case  of  the  Ethiopian;  and  that 
there  are  but  two  families  absolutely  confined  to  it, 
although  a  third  is  very  nearly  so.  While  sharing 
with  the  Ethiopian  region  the  want  of  several  groups  of  insectivores 
and  rodents,  such  as  the  typical  shrews  (Sorex),  marmots  (Arcto- 
mys),  and  voles  (Microtus),  it  lacks  some  of  the  other  deficiencies 
of  that  region,  true  pigs  (Sus)  and  deer  being  abundant,  although 
the  latter  belong  to  groups  distinct  from  those  of  the  Holarctic 
region,  while  bears,  belonging  to  two  genera,  are  likewise  met 


268  THE    ORIENTAL   REGION.  [CHAP. 


with.  Wart-hogs  (Phacochcerus),  aard-varks  (Orycteropus\  hyraces 
(Procaviid(B\  and  jumping-shrews  (Macroscelididce)  are  among 
some  of  the  more  characteristic  Ethiopian  animals  which  are 
wanting  in  the  Oriental  region  at  the  present  day,  and  have  not 
hitherto  been  obtained  there  in  a  fossil  state.  Giraffes,  a  number 
of  genera  of  antelopes,  and  hippopotami,  are  equally  conspicuous 
by  their  absence,  although  this,  as  we  have  seen,  is  but  a  com- 
paratively recent  feature  of  the  region,  since  most  of  these  forms 
are  represented  in  the  Pliocene  of  India  and  Burma.  A  notable 
feature  of  the  Oriental  as  distinct  from  the  Ethiopian  region  is  the 
circumstance  that  the  great  majority  of  its  fruit-bats  (like  those  of 
Madagascar)  belong  to  the  typical  genus  Pteropus,  which  is 
wanting  in  Ethiopia,  while  the  three  genera  of  the  family  found  in 
the  latter  area  are  absent  from  the  present  region.  Indeed  there 
is  a  very  curious  dissimilarity  between  the  flying-mammals  of  the 
two  areas,  Ethiopia  possessing  the  flying-squirrels  of  the  family 
Anomaluridce,  while  the  Oriental  flying-squirrels  all  belong  to  the 
SriuridiE,  the  genus  Pteromys  being  peculiar  to  the  region.  Among 
the  Insectivora,  the  so-called  flying -lemur  (Galeopithecus)  is  a 
peculiar  Oriental  type  which  has  no  Ethiopian  representative.  A 
somewhat  similar  instance  to  that  of  the  flying-squirrels  is  afforded 
by  the  tree-shrews  (Tupaiid&)  of  the  Oriental  region,  which  are 
represented  in  Ethiopia  by  the  jumping-shrews  (Macroscelidida). 
From  the  Holarctic  region,  the  Oriental  is  distinctly  differentiated 
by  the  presence  of  apes  and  lemurs  and  the  abundance  of 
monkeys,  together  with  the  presence  of  the  groups  mentioned 
above  as  being  now  restricted  to  this  and  the  Ethiopian  region, 
and  likewise  by  the  absence  of  the  typical  elaphine  deer, 
marmots,  susliks,  voles,  etc.,  and  the  scarcity  of  sheep  and  true 
goats,  which,  indeed,  enter  the  region  only  in  the  north-western 
frontier  of  India. 

Commencing  with  the  man-like  apes  of  the  family  Simiidce,  we 
find  the  Oriental  region  destitute  of  chimpanzees  and  gorillas, 
whose  place  is  taken  by  the  orangs  (Simla]  of  Borneo  and 
Sumatra,  characterised  by  the  reddish,  instead  of  blackish,  colora- 
tion of  their  hair,  and  their  more  wide  departure  from  the  human 
type.  Orangs  appear,  however,  to  have  inhabited  northern  India 
during  the  Pliocene,  and  as  chimpanzees  were  then  also  in  exist- 


VIII.] 


PRIMATES. 


269 


ence  there,  it  would  seem  that  the  region  must  be  regarded  as  the 
original  home  of  the  larger  man-like  apes.  The  smaller  long- 
armed  apes  known  as  gibbons  (Hylobates)  are  likewise  characteristic 
of  the  Oriental  region,  where  they  range  from  Assam  through  the 
Burmese  and  Malayan  countries  to  Hainan  and  Cambodia.  In 
the  upper  Miocene  these  apes  occurred  in  Central  Europe,  but — 
perhaps  on  account  of  their  small  size — their  remains  have  not 
hitherto  been  obtained  from  the  Indian  Pliocene.  Among  the 
Cercopithecidcz  the  Oriental  genera  of  monkeys  are  now  entirely 
distinct  from  those  of  Ethiopia,  although,  as  we  have  seen,  the 


FIG.  60.     SLOW  LORIS  (Nycticebus  tardigradus). 

African  genus  Papio  occurs  in  the  Indian  Pliocene  and  in  Madras 
survived  till  the  Plistocene.  And  here  it  may  be  noticed  that  this 
is  the  only  one  of  the  Ethiopian  genera  of  monkeys  that  is  found 
in  Arabia.  Of  the  other  genera,  Macacus  is  mainly  Oriental, 
although  with  representatives  in  northern  Africa,  Kashmir,  Tibet, 
and  Japan ;  while  the  langurs  (Semnopithecus]  are  likewise  almost 
wholly  confined  to  this  region,  although  they  range  into  Kashmir 
and  eastern  Tibet.  As  both  these  genera  are  found  in  the 
European  and  Indian  Pliocene,  they  are  evidently  ancient  types 
which  were  formerly  widely  spread  in  the  eastern  half  of  the  Old 


2/0  THE   ORIENTAL   REGION.  [CHAP. 

World.  Each  represents  a  sub-family  by  itself;  and  as  both  these 
sub-families  have  Ethiopian  genera,  which  are  unknown  in  a  fossil 
state,  it  may  be  suggested  that  the  latter  (like  the  wart-hogs)  have 
been  evolved  within  the  limits  of  the  Ethiopian  region  from  the 
Oriental  types.  To  the  same  sub-family  as  the  langurs  belongs  the 
singular  proboscis  monkey  (Nasalis)  of  Borneo.  Among  the 
lemuroid  Primates  there  are  Oriental  representatives  of  two 
families.  Of  these  the  Lemurida  include  the  lorises  of  the  genera 
Nycticebus  and  Loris,  the  former  ranging  over  the  Burmese, 
Malayan  and  Indo-Chinese  sub-regions,  while  the  latter  is  con- 
fined to  southern  India  and  Ceylon.  Although  these  animals  are 
nearly  allied  to  the  pottos  (Perodicticus)  of  western  Africa,  nothing 
is  known  as  to  their  past  history.  The  tarsiers  (Tarsiidce\  of 
which  there  is  but  a  single  genus  (Tarsius\  although  several 
specific  forms  have  been  recognised,  are  almost  confined  to  the 
Malayan  sub-region,  but  are  represented  in  Celebes,  as  they  are  in 
the  Philippines. 

Of  the  Insectivora,  the  most  aberrant  and  remarkable  forms  are 
the  flying-lemurs  (Galeopithecus),  constituting  a  sub-order  by  them- 
selves, and  ranging  from  south  Tenasserim  through  the  Malay 
peninsula  and  islands  to  the  Philippines.  As  with  the  tarsiers,  we 
have  no  palaeontological  history  of  these  creatures,  which  are 
probably  comparatively  modern  types.  The  most  characteristic 
Oriental  family  of  this  group  is  that  of  the  tree-shrews  (Tupaiidce], 
whose  members  have  the  form  and  habits  of  squirrels,  with  the 
structure  of  shrews.  The  typical  genus  Tupaia '  ranges  from  India 
throughout  the  Burmese  and  Malayan  regions,  but  is  unknown  in 
Ceylon ;  while  the  single  representative  of  the  pen-tailed  tree- 
shrews  (Ptilocercus],  characterised  by  the  pen-like  extremity  of  the 
exceedingly  long  tail,  is  confined  to  Borneo  and  some  of  the  adja- 
cent islands.  As  mentioned  in  an  earlier  chapter,  the  European 
Miocene  genera  Lanthanotherium  and  Galerix  appear  to  be  an- 
cestral types  of  this  family  ;  and  this  distribution  of  the  family  is  a 
well-marked  instance  of  the  curious  affinity  existing  between 
certain  mammalian  genera  of  the  middle  Tertiaries  of  Europe  and 

1  Two  species  (T.  murina  and  7".  frenata)  are  often  separated  as  Dendro- 
gale,  but  as  there  is  an  annectent  form  (see  Thomas,  Proc.  Zool.  Soc.  1892,  p. 
-225),  this  appears  unnecessary. 


VIII.] 


INSECTIVORA. 


2/1 


those  of  the  Malayan  sub-region,  the  absence  of  Tupaia  from 
Ceylon  probably  indicating  that  the  genus  is  essentially  a  Malayan 
one  which  has  immigrated  but  recently  into  India.  The  hedgehog 
family  (JErwaceidce),  which,  as  already  shown  is  an  ancient  one, 
has  a  very  remarkable  distribution  in  the  Oriental  region  ;  true 
hedgehogs  (Erinaceus]  ranging  into  India,  but  apparently  not  oc- 
curring in  Ceylon,  and  being  unknown  to  the  west  of  the  Bay  of 
Bengal.  In  the  latter  districts  their  place  is  taken  by  the  spineless 
and  more  rat-like  animals  forming  the  genus  Gymnura  and  the 


FIG.  61.     TREE-SHREW  {Tupaia  tana). 

allied  Hylomys  ;  and  here,  again,  we  have  to  note  the  occurrence 
of  an  allied  type  in  the  European  Oligocene,  which  has  been 
described  under  the  name  of  Necrogymnurus.  In  passing,  it  may 
be  remarked  that  the  survival  of  these  early  Tertiary  insectivorous 
types  in  the  Malayan  sub-region  serves  to  lend  support  to  the 
suggestion  made  in  a  previous  chapter1  that  opossums  may  also 
have  survived  in  the  same  area  long  after  they  had  ceased  to  exist 
in  western  Europe. 

Passing  over  the  moles  (Talpida),  of  which  the  typical  genus 
Talpa  only  just  impinges  on  the  region  in  the  frontiers  of  India> 

1  Supra,  pp.  51,  57- 


2/2  THE   ORIENTAL   REGION.  [CHAP. 

we  come  to  the  Soricidce  or  shrews.  Here  the  typical  shrews 
(Sorex)  are  wanting,  while  the  section  of  the  family  to  which  that 
genus  belongs  (characterised  by  the  reddish-brown  tips  to  the 
teeth)  is  represented  only  by  the  genus  Soriculus,  ranging  from  the 
southern  slopes  of  the  Himalaya  to  China.  Of  the  widely-spread 
musk-shrews  (Croddura)  it  is  unnecessary  to  speak;  but  it  may 
be  mentioned  that  of  the  two  almost  tailless  and  scaly-footed 
species  forming  the  genus  Annrosorex  one  is  from  Assam  and  the 
other  from  eastern  Tibet  and  Pekin.  Chimarrogale  includes  two 
aquatic  shrews,  one  of  which  is  found  in  the  eastern  Himalaya, 
the  hills  north  of  Burma,  and  M4  Kina  Balu  in  Borneo,  while 
the  second  is  Japanese. 

Among  the  Carnivora  the  region  is  especially  rich  in  Felidce, 
containing  more  species  than  any  other  part  of  the  world.  The 
tiger  (Felis  tigris)  is  usually  regarded  as  one  of  the  most  character- 
istic mammals  of  India,  but  as  its  range  extends  northwards  to 
Siberia,  while  its  fossilised  remains  have  been  found  within  the 
Arctic  circle,  and  it  is  unknown  in  Ceylon,  there  is  a  great  proba- 
bility that  this  feline  is  a  comparatively  recent  immigrant  into 
India  from  the  north-east.  The  range  of  the  lion  (F.  leo)  in  this 
•region  is  limited  to  India,  not  extending  to  the  eastward  of  the 
Bay  of  Bengal,  and  as  this  animal  was  widely  distributed  during 
the  Plistocene  in  Europe,  while  it  ranges  all  over  Africa,  it  may 
be  regarded  as  essentially  a  western  type,  or  exactly  the  opposite 
of  the  tiger.  Possibly  certain  remains  from  the  Indian  Plistocene 
which  have  been  assigned  to  the  latter  animal  may  really  belong  to 
the  former.  As  noticed  on  p.  234,  there  are  other  species  of 
Felis,  as  well  as  the  hunting-leopard  (CynteJurus),  which  are  com- 
mon to  India  and  Africa,  some  of  these  occurring  in  the  European 
Plistocene,  while  only  the  jungle-cat  (F.  chaus)  is  found  to  the 
eastward  of  the  Bay  of  Bengal,  and  there  not  further  east  than 
Burma.  On  the  other  hand,  there  are  certain  species,  like  the 
clouded  leopard  (F.  nebulosa)  and  the  marbled  cat  (F.  marmorata), 
which  are  essentially  eastern  forms,  their  range  including  the 
Malayan  sub-region  and  India,  but  not  Ceylon.  The  rusty- 
•spotted  cat  (F.  rubiginosa)  and  the  Indian  desert-cat  (F.  ornatd] 
are  species  whose  range  is  limited  in  one  case  to  India  and 
•Ceylon,  and  in  the  other  to  India  alone. 


VIII.]  CARNIVORA.  2/3 

In  the  civets  and  their  allies  ( Viverridtz]  the  Oriental  region 
approaches  the  Ethiopian  in  richness,  and  thereby  stands  in 
marked  contrast  to  the  Holarctic,  which  contains  only  a  single 
species  of  Genetta  and  another  of  Herpestes  in  southern  Europe, 
although  the  latter  genus  ranges  into  Kashmir.  Of  the  true  civets 
( Viverrd]  the  whole  of  the  species,  with  the  exception  of  one  from 
the  Ethiopian  region,  are  Oriental,  and  some  are  confined  to  the 
countries  to  the  east  of  the  Bay  of  Bengal;  one  small  species  being 
separated  by  many  zoologists  as  Viverricula.  The  beautifully- 
coloured  linsangs  (Linsangd)  are  exclusively  Oriental,  and  are  con- 
fined to  the  eastern  Himalayan  and  Malayan  sub-regions,  although 
represented  in  West  Africa  by  the  nearly  allied  Poiana.  Equally 
characteristic  of  the  region  are  the  two  species  of  Hemigale,  which 
are,  however,  exclusively  Malayan,  H.  hosei  being  limited  to  the 
mountains  of  North  Borneo.  The  palm-civets  of  the  genus  Para- 
doxurus  range  throughout  the  region,  and  have  also  representatives 
in  Celebes  :  their  place  being  taken  in  the  Ethiopian  region  by 
the  allied  genus  Nandinia.  On  the  other  hand,  the  two  species 
of  small-toothed  palm-civets  constituting  the  genus  Arctogale  are 
restricted  to  the  Burmese  and  Malayan  sub-regions  ;  the  same 
being  also  the  case  with  the  binturong,  which  is  the  sole  repre- 
sentative of  the  genus  Arctitis,  distinguished  by  its  pencilled  ears 
and  prehensile  tail.  Still  more  circumscribed  is  the  range  of  the 
peculiar  genus  Cynogale,  of  which  the  single  species  is  confined  to 
the  Malayan  sub-region.  All  the  foregoing  forms  belong  to  the 
sub-family  Viverrina  \  the  Herpestincz,  which  are  so  numerous  in 
the  Ethiopian  region,  being  represented  only  by  species  of  the 
large  and  widely-spread  genus  Herpestes.  Both  this  genus  and 
Viverra  date  from  the  European  Oligocene,  the  latter  also 
occurring  in  the  Pliocene  of  France  and  India ;  but  none  of  the 
others  are  known  in  a  fossil  state.  It  is,  however,  probable  that 
most  of  the  other  genera  are  comparatively  modern  derivatives 
from  the  original  stock ;  and  the  high  development  in  the  Malayan 
sub-region  of  a  group  first  known  from  the  Oligocene  of  Europe 
is  another  instance  of  the  relationship  of  the  faunas  of  these 
countries. 

Although  the  striped  hyaena  (Hycena  striata)  is  by  no  means 
confined  to  India,  its  range  extending  through  south-western  Asia 
L.  18 


274  THE   ORIENTAL   REGION.  [CHAP. 

to  northern  Africa,  it  is  unknown  in  Ceylon  or  in  the  countries  on 
the  eastern  side  of  the  Bay  of  Bengal,  which  forms,  indeed,  the 
present  limits  of  the  range  of  the  genus  in  this  direction.  As 
remains  of  the  existing  African  spotted  hyaena  (H.  crocutd)  have 
been  met  with  in  a  cave  in  Madras,  while  they  are  common  in  the 
Plistocene  of  southern  and  central  Europe,  it  is  manifest  that  both 
these  animals  are  as  essentially  western  types  as  is  the  lion.  And 
it  is  a  curious  circumstance  that  nearly  all  these  western  types  of 
mammals  ranging  into  India  (of  which  a  list  is  given  in  the  sequel) 
belong  to  genera  which  date  only  from  the  Miocene  or  Pliocene, 
whereas  very  many  of  the  Malayan  or  eastern  types  date  from  the 
Oligocene.  During  the  Pliocene  a  single  species  of  hyaena  ranged 
as  far  eastwards  as  China,  and  species  were  exceedingly  abundant 
in  India  at  the  same  epoch. 

As  regards  its  Canidcz,  the  Oriental  region  is  inferior  to 
the  Ethiopian  in  lacking  any  peculiar  generic  type,  although  it 
possesses  a  true  wolf  (Cants  pallipes),  and  three  species  of  wild 
dog  (C.  rutilans,  etc.),  the  latter,  on  account  of  the  absence  of  the 
last  tooth  in  the  lower  jaw  and  other  differences,  being  frequently 
referred  to  a  distinct  genus,  under  the  title  of  Cyan.  Whereas, 
however,  the  Indian  wolf,  which  ranges  into  southern  Arabia,  is 
unknown  either  in  Ceylon  or  in  the  countries  to  the  east  of  the 
Bay  of  Bengal,  the  wild  dogs  are  found  throughout  the  region,  and 
have  also  a  representative  beyond  it  in  the  mountains  of  Central 
Asia,  and  they  are  likewise  known  by  fossil  species  from  the 
European  Plistocene.  The  wolf,  which  is  very  closely  allied  to 
the  European  species,  may  be  the  descendant  of  a  fossil  form 
found  in  the  Siwaliks,  but  its  absence  from  Ceylon  would  seem  to 
indicate  that  it  has  only  reached  southern  India  at  a  comparatively 
modern  date.  No  foxes  are  known  to  the  east  of  the  Bay  of 
Bengal,  and  the  jackal  does  not  range  east  of  Burma. 

The  Oriental  region  is  the  home  of  three  well-marked  species 
of  bears,  and  thereby  presents  a  decided  contrast  to  the  Ethiopian. 
Of  these  the  Himalayan  black  bear  (Ursus  torquatus)  ranges  from 
the  forest  districts  of  the  Himalaya  to  Burma,  and  thence  to  the 
Indo-Chinese  sub-region.  The  small  Malayan  bear  (U.  malay- 
anus]  is  restricted  to  the  Burmese  and  Malayan  sub-regions  ;  and 
the  great  Indian  sloth  bear  (Melursus  ursinus],  which  is  the  sole 


VIII.] 


CARNIVORA. 


275 


representative  of  a  separate  genus,  is  confined  to  India  and 
Ceylon,  and  is  known  to  have  been  an  inhabitant  of  Madras  since 
the  Plistocene  era.  Not  improbably  it  may  be  the  descendant  of 
a  Siwalik  species  ( U.  theobaldi),  which  is  the  earliest  known  repre- 
sentative of  the  true  bears;  and  the  Malayan  species  may  be 


FIG.  62.     INDIAN  SLOTH-BEAR  (Melursus  ursinus). 

derived  from  a  small  extinct  bear  whose  remains  occur  in  the 
Plistocene  of  the  Narbada  valley. 

One  of  the  most  remarkable  of  Oriental  carnivores  is  the 
panda,  or  cat-bear  (sElurus  fulgens),  which  ranges  from  the  East- 
ern Himalaya  to  Yunnan,  and  is  the  single  existing  Old  World 
representative  of  the  Procyonida.  Curiously  enough,  the  remains 
of  a  much  larger  species  of  the  same  genus  have  been  discovered 
in  the  English  Pliocene ;  and  it  is  thus  evident  that  sElurus 
formerly  enjoyed  a  wide  range.  From  the  restriction  of  all  the 
other  known  members  of  the  family  to  the  New  World,  fossil 
types  may  be  looked  for  in  eastern  Asia,  as  it  is  quite  clear  that 
the  distributional  area  of  the  group  must  once  have  been  con- 
tinuous. 

In  the  Mustelidce  there  are  four  generic  types  very  character- 

18— 2 


276 


THE   ORIENTAL   REGION. 


[CHAP. 


istic  of  the  region,  although  two  of  them  are  not  confined  to  it. 
The  first  of  these  comprises  the  three  species  of  sand-badger 
(Arctonyx),  two  of  which  are  found  in  the  Himalayan  and  Burmese 
sub-regions,  while  the  third  is  Tibetan.  The  single  Oriental 
species  of  ratel  (Mellivora)  is  restricted  to  India,  exclusive  of 
Ceylon ;  a  fossil  species  occurring  in  the  Siwaliks.  The  only  other 
living  form  is  Ethiopian.  Its  distribution  would  thus  seem  to 
indicate  that  the  genus  originated  in  northern  India,  whence  it 


FIG.  63.     INDIAN  RATEL  (Mellivora  ratel). 

migrated  into  Africa  while  there  was  a  free  communication  between 
the  two  continents,  and  that  it  only  reached  southern  India  (where 
it  is  unknown  on  the  Malabar  coast)  at  a  comparatively  recent 
epoch.  The  third  genus,  Helictis,  comprising  badger-like  animals 
with  long  bushy  tails,  is  represented  by  four  species,  ranging  from 
India  to  China,  but  unknown  in  Ceylon.  Lastly,  the  teledu,  or 
small  burrowing  badger  (Mydaus  meliceps)  of  the  Malayan  sub- 
region,  is  the  sole  representative  of  its  genus,  and  is  found  at  con- 


VIII.]  RODENTIA.  277 

siderable  elevations  in  Java,  as  well  as  in  Sumatra  and  Borneo. 
No  fossil  representatives  of  either  of  these  two  genera  are  at 
present  known. 

Among  the  rodents,  the  grooved-toothed  squirrel  (Rhithro- 
scturus)  is  a  peculiar  type  confined  to  the  island  of  Borneo; 
and  the  pigmy  squirrels  (Nannosciurus)  are  represented  by  four 
Malayan  species,  the  only  other  member  of  the  genus  being  West 
African.  The  true  squirrels  (Sciurus),  as  mentioned  above,  attain 
their  maximum  development  in  the  Malayan  sub-region.  The 
flying  -  squirrels  are  represented  by  the  genera  Pteromys  and 
Sciuropterus,  the  former  being  exclusively  Oriental,  and  the  latter 
having  one  species  in  the  eastern,  and  a  second  in  the  western 
half  of  the  Holarctic  region,  in  addition  to  being  represented  in  a 
fossil  state  in  the  French  Miocene.  In  the  Muridcz  there  are  no 
less  than  eleven  genera — in  most  cases  respectively  represented 
by  only  a  single  species — peculiar  to  this  region,  while  another  is 
Oriental  and  Ethiopian  only.  Of  the  peculiar  types,  one  of  the 
most  remarkable  is  Chrotomys,  from  the  mountains  of  Luzon,  in 
the  Philippines,  belonging  to  the  sub-family  Hydromyince,  of  which 
the  typical  forms  are  Australian1.  From  other  members  of  the 
sub-family  the  single  species  of  this  genus  differs  in  having  three 
(in  place  of  only  two)  pairs  of  molar  teeth,  thereby  forming  a 
link  with  ordinary  murines.  This  animal,  which  is  about  the 
size  of  a  rat,  is  easily  recognised  by  the  presence  of  an  orange  or 
buff  line  running  down  the  middle  of  the  back.  Luzon  has  also 
yielded  another  rat,  provisionally  referred  to  the  Australian  genus 
Xeromys*.  Another  unique  Oriental  type  is  found  in  the  long- 
tailed  dormouse-like  form  from  the  Malabar  coast  known  as 
Platacanthomys,  which  constitutes  a  sub-family  by  itself.  Phlao- 
mys,  likewise  representing  a  separate  group  of  the  same  rank,  is 
restricted  to  the  Philippines,  and  is  characterised  by  the  molar 
teeth  being  divided  into  three  transverse  lobes.  The  one  species 
is  of  very  large  size.  Nearly  allied  is  a  huge,  rough-haired,  grey 
or  blackish  rat,  from  the  mountains  of  Luzon,  which  may  be 
compared  in  size  to  a  small  marmot,  and  for  which  the  name 
Crateromys  has  been  suggested.  This  differs  from  Phlceomys  by 


1  Vide  suprh,  p.  40. 

2  Appendix,  No.  31 


2/8  THE   ORIENTAL   REGION.  [CHAP. 

the  smaller  claws  and  more  bushy  tail,  and  also  by  the  completely 
tuberculate  molars.  The  single  species  of  the  Burmese  rat-like 
Hapalomys  differs  from  all  other  members  of  the  sub-family 
Murincz  in  possessing  three  longitudinal  rows  of  tubercles  on 
the  lower  as  well  as  on  the  upper  molar  teeth.  The  one  repre- 
sentative of  the  allied  genus  Vandeleuria  ranges  from  India  and 
Ceylon  to  Yunnan.  The  pencil-tailed  tree-mice  (Chiropodomys), 
of  which  there  are  three  species,  are  restricted  to  the  Burmese  and 
Malayan  sub-regions;  and  the  small  red  rat  representing  the  genus 
Pithechirus  is  known  only  from  Sumatra  and  Java.  With  the 
shrew-rat  (Rhynchomys]  we  revert  to  several  peculiar  forms  from 
the  mountains  of  Luzon,  in  the  Philippines.  This  rodent,  which 
is  about  the  size  of  an  ordinary  rat,  has  the  muzzle  extraordinarily 
slender  and  elongated,  with  very  feeble  incisors,  and  it  is  pro- 
bable that  it  lives  on  animal  substances,  possibly  caterpillars. 
The  two  other  Philippine  types  form  the  genera  Carpomys  and 
Batomys ;  the  former  with  two,  and  the  latter  with  a  single 
species.  They  are  more  or  less  dormouse-like  forms,  with  blunt 
muzzles,  thick  woolly  fur.  and  long  and  well-haired  tails.  Lastly, 
the  bush-rats  (Golunda)  have  one  Indian  and  another  Ethiopian 
representative. 

An  interesting  instance  of  how  the  present  distribution  of  a 
genus  is  explained  by  palaeontological  discoveries  is  afforded  by 
the  brush-tailed  porcupines  (Atherura],  now  represented  by  one 
species  from  the  Malayan,  and  a  second  from  the  West  African 
sub-region ;  the  connecting  form  being  one  of  which  fossil  teeth 
have  been  found  in  the  Karnul  district  of  Madras.  From  this 
it  may  be  inferred  that  the  genus  was  probably  also  represented 
in  the  Siwaliks.  To  the  same  family  (Hystricida)  belongs  a 
peculiar  porcupine  from  Borneo,  constituting  the  genus  Trichys. 

Passing  on  to  the  ungulates,  we  have  first  to  notice  a  peculiar 
group  of  oxen  forming  a  section  of  the  genus  Bos,  which  is  con- 
fined to  this  region,  and  characterised,  in  addition  to  certain  features 
of  the  skull  and  horns,  by  the  dark  colour  of  the  males,  or  of 
both  males  and  females.  Of  these,  the  gaur  (B.  gaurus)  inhabits 
both  India  and  the  Malayan  countries,  but  appears  never  to  have 
reached  Ceylon  ;  while  the  banteng  (B.  sondaicus)  is  confined  to 
the  countries  on  the  east  of  the  Bay  of  Bengal.  Fossil  representa- 


VIIL] 


UNGULATA. 


2/9 


tives  of  this  group  occur  in  the  Indian  Plistocene  ;  and  certain 
generalised  oxen  from  the  Siwalik  Hills  and  the  Pliocene  of 
southern  Europe,  in  which  the  females  were  generally  or  always 
hornless,  may  have  been  the  ancestral  type.  The  Indian  buffalo 
(B.  bubalus]  is  markedly  distinct  from  the  Ethiopian  forms,  and 
has  ancestral  representatives  in  the  Indian  Pliocene  and  Plisto- 
cene. While  abundant  in  Ceylon,  it  is  probably  unknown  in  a 
truly  wild  state  to  the  east  of  the  Bay  of  Bengal.  The  Philippine 
buffalo,  or  tamarao  (B.  mindorensis)  is  regarded  by  some  as  a 


FIG.  64.    JAPANESE  SEROW  (Nemorhadiis  crispus}. 

cross  between  the  last  and  the  anoa  of  Celebes ;  ancestral  types  of 
the  latter  occurring,  as  already  mentioned,  in  the  Siwaliks.  In  the 
same  family  the  short-horned  goats  of  the  genus  Hemitragus  are 
represented  by  two  Indian  species,  one  inhabiting  the  Himalaya 
and  the  other  the  Nilgiris ;  the  third  living  species  being  south 
Arabian.  One  extinct  species  occurs  in  the  Siwaliks  and  a  second 
one  in  Perim  Island,  so  that  the  group  is  essentially  an  Indian 
one ;  and,  as  already  mentioned,  its  present  distribution  can  only 


280  THE   ORIENTAL   REGION.  [CHAP. 

be  accounted  for  by  a  lowering  of  the  temperature  during  a  past 
epoch.  Of  the  goat-like  genera  Nemorhcedtis  and  Cemas,  the 
former  has  a  wide  range  in  the  region  and  also  extends  into 
northern  China  and  Japan,  while  the  latter  is  represented  solely 
by  the  Himalayan  goral ;  no  fossil  types  of  either  being  known. 
In  its  poverty  of  antelopes  (exclusive  of  the  widely-spread  gazelles) 
the  Oriental  presents  a  most  remarkable  contrast  to  the  Ethiopian 
region,  although  this  poverty  is  largely  a  feature  of  the  present 
epoch,  African  types  being  common  in  the  Siwaliks.  The  sole 
existing  forms  are  the  four-horned  antelope  (Tetraceros  quadri- 
cornis],  the  black-buck  (Antilope  cervicapra)^  and  the  nilgai  (Bos- 
elaphus  tragocamelus],  each  of  which  forms  a  genus  by  itself,  and 
all  of  which  are  restricted  to  India,  exclusive  of  Ceylon.  Indeed, 
it  is  a  remarkable  feature  that  true  antelopes  and  gazelles  are 
unknown  to  the  eastward  of  the  Bay  of  Bengal ;  although  this  may 
be  chiefly  or  entirely  due  to  the  countries  to  the  eastward  being 
unsuited  to  their  habits.  The  nilgai,  which  has  fossil  representa- 
tives in  the  Indian  Plistocene  and  Pliocene,  is  allied  to  the  kudu 
group  of  Africa,  while  the  four-horned  antelope  is  a  near  relative 
of  the  duikers.  It  will  be  unnecessary  to  say  anything  with  regard 
to  the  true  goats  (Caprd)  and  sheep  (Ovis)  inhabiting  the  region, 
since  these  are  found  only  on  the  north-western  frontier  of  India, 
and  are  obviously  intruders  from  the  Holarctic  region.  It  is, 
however,  important  to  mention  that  extinct  representatives  of  one, 
if  not  of  both  groups,  occur  in  the  Siwalik  Hills. 

The  abundance  of  Cervidce  is  one  of  the  most  noticeable 
features  distinguishing  the  Oriental  from  the  Ethiopian  region ; 
there  being  an  equally  marked  difference  in  this  respect  between 
the  former  and  the  Holarctic  area.  Although  the  majority  of  the 
Oriental  deer  are  now  included  in  the  genus  Cervus,  the  typical, 
or  elaphine  group,  as  represented  by  the  red  deer  and  the  wapiti, 
is  entirely  wanting,  its  place  being  taken  by  the  sambur  and  its 
allies  (C.  unicolor),  forming  the  rusine  group;  the  swamp-deer 
(C.  duvaucelt),  which  with  another  species  constitutes  the  rucervine 
group,  and  the  Indian  spotted  deer  (C.  axis),  alone  representing 
the  axine  group.  Rusine  deer  are  abundant  in  the  Indian 
Siwaliks,  but  appear  to  be  unknown  in  the  Pikermi  beds. 
Although  they  have  one  Tibetan  representative,  the  smaller  deer 


VIII.]  UNGULATA.  28 1 

known  as  muntjacs  (Cervulus) — which  are  characterised  by  the 
length  of  the  pedicles  of  the  antlers  and  the  shortness  of  the  antlers 
themselves — form  a  very  characteristic  Oriental  group,  ranging  over 
the  entire  region.  Not  improbably  they  are  represented  in  the 
Pliocene  of  Europe. 

The  chevrotains,  or  TragulidcK^  which  have  already  been  shown 
to  be  abundant  in  the  European  Oligocene  and  Miocene — remains 
of  the  West  African  genus  occurring  in  the  latter  deposits  and 
the  Indian  Pliocene — are  represented  in  the  Oriental  region  by 
Tragulus,  which  dates  from  the  Siwalik  epoch,  and  ranges  from 
India  and  Ceylon  to  the  Philippines.  Although  wild  camels  are 
now  everywhere  unknown,  it  is  probable  that  India  and  the 
Holarctic  region  was  their  original  home,  remains  of  the  genus 
Camelus  being  found  in  the  Pliocene  of  the  Siwalik  Hills. 

The  large  number  of  species  of  true  pigs  (Sus}  characterising 
the  Oriental  region  is  a  notable  feature,  India  itself  being  in- 
habited by  a  species  (Sus  cristatus]  nearly  allied  to  the  European 
wild  boar,  while  the  Malayan  sub-region  is  the  home  of  a  consider- 
able number  of  species  differing  more  or  less  markedly  from  the 
latter.  The  genus  is  well  represented  both  in  the  Pliocene  and 
Plistocene  of  India,  but  in  neither  of  these  formations  are  there 
any  of  the  Ethiopian  types  of  the  family. 

With  the  exception  of  the  Ethiopian,  the  Oriental  region  is  now 
the  sole  one  where  the  family  Rhino cerotida  still  exists ;  but  there 
is  a  remarkable  difference  between  the  species  inhabiting  the  two 
areas,  all  the  three  living  Asiatic  forms  being  furnished  with  teeth 
in  the  front  of  the  jaws,  which,  as  we  have  seen  in  the  last  chapter, 
are  wanting  in  the  African  species.  While  one  of  the  Oriental 
rhinoceroses  (R.  sondaicus}  ranges  from  eastern  Bengal  to  the 
Malayan  islands,  and  a  second  (R.  sumatrensis)  from  Assam  to 
the  same,  the  great  Indian  species  (R.  unicornis)  is  unknown  to 
the  eastward  of  Assam,  as  it  is  in  Ceylon.  Fossil  remains  of  the 
latter  are  found  in  the  Plistocene  of  the  Narbada  valley,  while 
ancestral  types  both  of  this  species  and  of  R.  sondaicus  are  met 
with  in  the  Pliocene  of  the  Siwalik  Hills.  It  is,  however,  very 
remarkable  that  Ethiopian  types  of  the  genus  occur  not  only  in 
the  last-named  deposits,  but  likewise  in  the  Plistocene  of  Madras  ; 
the  total  extinction  of  this  group  in  India  being,  as  in  the  case  of 


282  THE   ORIENTAL   REGION.  [CHAP. 

other  Ethiopian  types,  almost  impossible  to  account  for.  One  of 
the  two-horned  extinct  Indian  rhinoceroses  (R.  platyrhinus]  ap- 
pears to  have  been  the  ancestor  both  of  the  existing  R.  simus  of 
Africa  and  R.  antiquitatis  of  the  Plistocene  of  northern  Asia  and 
Europe  ;  the  evolution  of  the  latter  species  having  not  improbably 
taken  place  in  the  countries  lying  between  India  and  China, 
whence  the  creature  wandered  northwards  and  westwards  with  the 
mammoth  to  the  Arctic  tundras.  With  regard  to  the  Equidce,  it 
will  suffice  to  mention  that  species  of  Equus  occur  in  the  Plisto- 
cene of  Central  India  and  Madras,  and  that  wild  asses  (of  a  type 
markedly  different  from  the  African  wild  ass)  occur  in  Sind  and 
Kach.  The  genus,  like  the  antelopes,  is,  however,  totally  un- 
known in  the  countries  to  the  east  of  the  Bay  of  Bengal,  as  it  is  in 
Ceylon.  In  the  Tapiridcz,  the  Malayan  tapir  (Tapirus  indicus) 
inhabits  the  Malay  Peninsula  as  far  north  as  Mergui,  and  also  the 
islands  of  Sumatra  and  Borneo.  It  is  important  to  notice  that 
although  fossil  remains  of  tapirs  are  unknown  from  the  Pliocene  of 
the  Siwaliks,  they  are  met  with  in  caverns  in  China. 

Distinguished,  among  other  features,  from  its  African  cousin  by 
the  thinner  and  more  numerous  enamel-plates  of  its  molar  teeth, 
the  Indian  elephant  (Elephas  indicus)  ranges  over  the  greater  part 
of  the  region,  being  found  in  suitable  districts  in  India,  Ceylon, 
Burma,  the  Malay  Peninsula,  Cochin  China,  and  Sumatra.  This 
species  is  a  near  ally  of  the  mammoth  (E.  primigenius) ;  and  it 
may  prove  that  both  are  descendants  of  a  Siwalik  species  (E. 
hysudricus],  which  has  molar  teeth  of  the  type  we  should  expect  to 
find  in  such  an  ancestral  form.  If  this  view  be  correct,  the 
mammoth  has  probably  wandered  to  northern  Europe  and  Siberia 
from  the  countries  lying  just  to  the  east  of  India.  It  has  been 
mentioned  in  an  earlier  chapter  that  the  extinct  so-called  stego- 
dont  elephants  (such  as  E.  clifti  and  E.  insignis]  are  mainly 
confined  to  this  region,  although  some  of  the  species  are  found  in 
north  China  and  Japan.  As  these  elephants  form  the  transition 
between  Elephas  and  Mastodon,  and  also  since  the  species  of  the 
latter  genus  which  may  be  regarded  as  the  original  stock  of  the 
elephants  is  confined  to  the  Indian  and  Malayan  Pliocene,  it  may 
be  taken  for  granted  that  the  elephants  have  been  developed  from 
the  mastodons  within  the  limits  of  the  Oriental  region.  In  the 


VIII.]  PANGOLINS.  283 

Plistocene  of  the  Narbada  Valley  in  central  India  there  occurs  a 
species  (E.  namadicus]  closely  allied  to  the  contemporary  Euro- 
pean E.  antiqims,  in  both  of  which  the  molars  are  intermediate  in 
structure  between  those  of  the  living  Indian  and  African  species. 
Elephas  planifrons  of  the  Siwaliks,  which  has  molars  of  a  still  more 
generalised  type,  is  equally  closely  allied  to  E.  meridionalis  of  the 
upper  Pliocene  of  Europe;  and  it  is  quite  probable  that  the 
former  may  be  the  original  ancestral  stock  of  the  African  elephant. 
It  is  worth  mentioning  that  the  stegodont  elephants  survived  till 
the  Plistocene ;  and  also  that  some  of  the  species  of  this  group 
inhabiting  India,  as  well  as  certain  mastodons,  ranged  as  far  east- 
wards as  Java,  Borneo,  China,  and  Japan. 


FlG.  65.     WHITE-BELLIED  PANGOLIN  (Manis  tricuspis). 

The  last  mammals  that  we  have  to  mention  are  the  pangolins 
(Mamdcz),  which  are  now  common  to  the  Oriental  and  Ethiopian 
regions,  and  appear  to  be  represented  by  an  extinct  genus  in  the 
European  upper  Oligocene.  The  presence  of  horny  scales  in- 
vesting the  whole  of  the  body  and  tail  serves  to  distinguish  the 


284  THE   ORIENTAL   REGION.  [CHAP. 

pangolins  from  all  other  mammals  whatsoever  ;  and  the  Oriental 
species  are  further  characterised  by  having  the  median  series  of 
scales  on  the  body  continued  to  the  tip  of  the  tail,  and  likewise  by 
the  presence  of  numerous  isolated  hairs  between  the  scales  of  the 
back,  as  well  as  by  the  presence  of  small  ears.  Of  the  three 
Oriental  species,  Manis  javanica  ranges  from  Burma  through  the 
Malay  Peninsula  to  Java  and  Borneo  ;  M.  aurita  extends  from 
Nipal  to  the  Indo-Chinese  sub-region  ;  while  M.  pentedactyla  is 
restricted  to  India  and  Ceylon.  The  most  remarkable  feature 
connected  with  the  distribution  of  the  group  is,  however,  the 
circumstance  that  claw-bones  indistinguishable  from  those  of  the 
giant  pangolin  (M.  giganted]  of  West  Africa  have  been  discovered 
in  a  cavern  in  the  Karnul  district  of  Madras. 

In  the  following  list  the  leading  results  of  the  foregoing  survey 
are  put  in  tabular  form,  the  italics  indicating  groups  or  species 
peculiar  to  the  region. 

I.     Primates. 


Simia.     Borneo  and  Sumatra  ;  fossil  in  India. 
Hylobates.     Burmese  and  Malayan  ;  fossil  in  European 
Miocene. 

CERCOPITHECID^E. 

Macacus.     Now    mainly  Oriental,  but   occurring   on 

the    southern  borders  of  the   Holarctic   region  ; 

fossil  in  the  European  and  Indian  Pliocene. 
Semnopithecus.     An  outlying  species  in  Eastern  Tibet 

and  one  in  Kashmir  ;  fossil  in  Pliocene  of  Europe 

and  India. 
Nasalis.     Borneo. 


Nycticebus.     Burmese,  Malayan,  and  Indo-Chinese. 
Loris.     S.  India  and  Ceylon. 


Elsewhere  only  in  Celebes. 
Tarsius.     Malayan,  extending  into  Celebes. 


VIII.]  LIST   OF   THE  FAUNA.  285 

II.  Insectivora. 

GALEOPITHECID&. 

Galeopithecus.     Malayan. 

TUPAIIDsE. 

Tupaia.     Indian  and  Malayan. 

Ptilocercus.     Borneo  and  some  adjacent  islands. 

ERINACEID^E. 

Gymnura.     Burmese  and  Malayan. 
Hylomys.     Burmese  and  Malayan. 

SORICID/E. 

Soriculus.     Himalayan  and  Indo-Chinese. 

Anurosorex.  Known  by  one  species  from  Assam  and 
a  second  from  Tibet  and  Pekin. 

Chimarrogale.  Represented  by  one  species  from  the 
eastern  Himalaya,  hills  north  of  Burma,  and  Mt 
Kina  Balu,  Borneo,  and  a  second  from  Japan. 

III.  Carnivora. 

FELID^:.     Very  numerous  in  the  region. 

Cynaelurus.  Indian  and  Ethiopian  ;  the  one  species 
being  common  to  the  two  areas  ;  fossil  in  Indian 
Pliocene. 

VlVERRID/E. 

Viverra.  All  the  species,  except  a  single  Ethiopian 
one,  are  Oriental,  one  of  these  being  frequently 
regarded  as  the  representative  of  a  distinct  genus 
( Viverricula) ;  fossil  in  European  Oligocene  and 
European  and  Indian  Pliocene. 

Hemigale.     Malayan. 

Linsanga.     Malayan  and  E.  Himalayan. 

Paradoxurus.     An  outlying  species  in  Celebes. 

Arctogale.     Burmese  and  Malayan. 

Arctictis.     Burmese  and  Malayan. 

Cynogale.     Malayan. 

URSID^E. 

Melursus.     India  and  Ceylon. 


286  THE   ORIENTAL   REGION.  [CHAP. 

III.  Carnivora.     (Cont.) 

PROCYONID^E. 

sElurus.     Eastern  Himalayan  and  Burmese ;  fossil  in 

English  Pliocene. 
MUSTELID^E. 

Arctonyx.     Two  E.  Himalayan  and  Burmese  species, 

and  probably  a  third  from  Tibet. 
Mellivora.     One     Indian     and     another     Ethiopian 

species  ;    fossil  in  Indian  Pliocene. 
Helictis.     India  to  China. 
Mydaus.     Malayan. 

IV.  Rodentia. 

SCIURID^E. 

Rhithrosciurus.     Borneo. 

Nannosciurus.     Represented   elsewhere    by    a    single 

West  African  species. 
Sciurus.     This  almost  cosmopolitan  genus  appears  to 

attain  its  maximum  development  in  the  Malayan 

sub-region. 
Pteromys. 
MURID^E. 

Chrotomys.     Philippines  (Luzon). 
Xeromys.     Philippines  (Luzon),  and  Australia. 
Phlceomys.     Philippines. 
Crateromys.     Philippines  (Luzon). 
Hapalomys.     Burma. 
Vandeleuria.     India  and  Burma. 
Chiropodomys.     Burmese  and  Malayan. 
Pithechirus.     Sumatra  and  Java. 
Rhynchomys.  \ 

Carpomys.      >•  Philippines  (Luzon). 
Batomys.       / 

Golunda.     One  Indian  and  one  Ethiopian  species. 
HYSTRICID^E. 

Atherura.     One    Malayan    and    one    West    African 

species ;    fossil  in  Indian  Plistocene. 
Trichys.     Borneo. 


VIII.]  LIST   OF   THE    FAUNA.  287 

V.     Ungulata. 


Bos.     The  Bibovine  group  exclusively  Oriental. 
Hemitragus.     Two  Indian  species,  and  a  third  in  the 

South  Arabian  sub-region  of  Ethiopia;  fossil  in 

Indian  Pliocene. 
Nemorhaedus.     Largely    Oriental    (Himalayan,    Bur- 

mese, and  Malayan),  but  extending  into  northern 

China  and  Japan. 
Cemas.     Himalayan. 
Tetraceros.     Indian. 

Antilope.     Indian  ;  fossil  in  Plistocene. 
Boselaphus.     Indian  ;  fossil  in  Plistocene  and  Pliocene. 


Cervus.  The  Rusine,  Ruce.rvine,  and  Axine  groups 
of  this  genus  are  characteristic  of  the"  region, 
although  the  first  is  also  represented  in  the  Austro- 
Malayan. 

Cervulus.  Mainly  Oriental,  but  with  one  Tibetan 
species. 

TRAGULID^E. 

Tragulus.  India,  Ceylon,  and  Malayan  sub-region  ; 
fossil  in  Indian  Pliocene. 


Sus.  Attains  its  maximum  specific  development  in 
the  Malayan  sub-region. 

RHINOCEROTID^E. 

Rhinoceros.  The  three  existing  Oriental  species 
differ  from  the  Ethiopian  forms  in  having  front 
teeth. 

TAPIRID^E. 

Tapirus.  Malayan  :  elsewhere  living  only  in  the 
Neogaeic  realm,  but  widely  distributed  in  a  fossil 
state,  although  absent  from  the  Siwaliks. 


288  THE   ORIENTAL   REGION.  [CHAP. 

V.  Ungulata  (cont.). 

ELEPHANTID^E. 

Elephas.  The  existing  Oriental  elephant  is  widely 
different  from  the  Ethiopian,  although  nearly 
allied  to  the  Holarctic  mammoth ;  the  extinct 
Stegodont  group  is  mainly  Oriental,  although 
extending  into  north  China  and  Japan. 

VI.  Effbdientia. 

MANID^E. 

Manis.  Elsewhere  only  in  Ethiopian  region ;  fossil 
in  Indian  Plistocene. 

The  relations  of  peninsular  India  to  the  Himalayan  area  have 
been  already  discussed  at  the  commencement  of 
of^hV  Regiony  this  cnaPter \  wm'le  tne  land-connection  which 
appears  to  have  existed  between  India  and  Mada- 
gascar, and  thus  with  Africa,  has  been  alluded  to  in  an  earlier 
one.  The  latter  connection  must  have  ceased  to  exist  before 
the  Pliocene  era ;  and,  as  we  have  seen,  the  descendants  of  the 
Siwalik  mammals  would  appear  to  have  made  their  way  into 
Ethiopia  across  Syria  or  Arabia.  During  the  Pliocene,  India,  at 
least,  could  not  have  been  distinguished  as  a  region  from  Ethiopia 
as  it  exists  at  the  present  day ;  and  even  in  the  Plistocene  the 
connection  between  the  faunas  of  the  two  areas  was  much  more 
intimate  than  it  is  now.  The  full  reason  for  this  gradual  dis- 
appearance of  the  modern  Ethiopian  types  from  the  Indian  area 
will  probably  never  be  known  ;  but  there  can  be  little  doubt  that 
the  gradual  refrigeration  of  the  northern  hemisphere  with  the 
advent  of  the  glacial  period  has  been  largely  instrumental ;  the 
present  distribution  of  Hemitragus  being  only  explicable  on  the 
hypothesis  of  a  marked  lowering  of  the  temperature  over  India. 

The  more  peculiar  mammals  now  inhabiting  the  Oriental 
region  may  be  roughly  arranged  under  five  headings.  The  first 
will  include  those  that  are  common  to  India  and  some  of  the 
countries  to  the  west  or  south-west,  but  are,  for  the  most  part, 
unknown  in  either  Ceylon  or  the  countries  to  the  eastward  of  the 
Bay  of  Bengal.  Under  this  category  may  be  included  the  follow- 
ing, viz. :— 


VIII.] 


LIST   OF   THE   FAUNA. 


289 


INSECTIVORA.     Erinaceus. 

CARNIVORA.       Felis    leo.       Ethiopian,    and    European    Plisto- 

cene. 
Felis  chaus.     Ethiopian,  and  European  Plistocene ; 

ranges  eastward  into  Burma. 
Felis  caracal.     Ethiopian. 
Cynaelurus  jubatus.     Ethiopian. 
Hyaena  striata.     Western  Asia  and  North  Africa. 
Hyaena  crocuta.     Ethiopian  ;  occurs  both  in  India 

and    Europe    during    the    Plistocene.     No 

representative  of  the  genus  known  in  the 

Burmese  or  Malayan  countries. 
Canis    aureus.     S.    W.    Asia ;    also    ranges    into 

Burma. 

Canis  pallipes.     South  Arabian. 
Mellivora.     The   Indian   and   Ethiopian    species 

very  closely  allied. 

RODENTIA.         Golunda.     Ethiopian  ;  the  Indian  species  ranges 
into  Ceylon. 

UNGULATA.        Hemitragus.     South  Arabian. 

Antelopes.  )  None   known   east   of   the    Bay   of 
Equus.         j          Bengal. 

The  second  group  includes  such  genera  and  species  as  are 
common  to  India  and  Ceylon,  but  are  unknown  elsewhere.  Here 
we  have : — 


PRIMATES. 
CARNIVORA. 

RODENTIA. 
UNGULATA. 


Loris. 

Felis  rubiginosa. 
Melursus. 

Golunda  ellioti. 


Bos  bubalus. 
Cervus  axis. 
Tragulus  memimna. 

EFFODIENTIA.    Manis  pentedactyla. 
L. 


290 


THE   ORIENTAL   REGION. 


[CHAP. 


The  third  group,  which  is  a  small  one,  comprises  types  which 
are  confined  to  India ;  it  includes 
Felis  ornata. 
Boselaphus. 
Tetraceros. 
Antilope. 
Rhinoceros  unicornis. 

In  the  fourth  group  we  have  generic  or  specific  forms  common 
to  India  and  the  countries  to  the  eastward  of  the  Bay  of  Bengal, 
but  unknown  in  Ceylon  or  in  the  countries  to  the  west  or  south- 
west.    This  list  comprises  the  following,  viz.  : — 
PRIMATES.          Hylobates. 
INSECTIVORA.     Tupaia. 
CARNIVORA.        Felis  tigris. 

Felis  nebulosa. 

Felis  marmorata. 

^Elurus. 

Helictis. 

Arctonyx. 

UNGULATA.        Bos  gaums. 

Bos  frontalis. 

Nemorhaedus. 

Cervus  porcinus. 

Finally,  we  have  (among  others)  the  following  group  of  genera 
and  species  confined  to  the  countries  lying  immediately  to  the 
eastward  of  the  Bay  of  Bengal,  viz.  :— 

PRIMATES.          Simia. 

Nasalis. 

Nycticebus. 

Tarsius. 
INSECTIVORA.     Galeopithecus. 

Gymnura. 

Hylomys. 

CARNIVORA.       Mydaus. 
RODENTIA.         Rhithrosciurus. 

Trichys. 


VIII.]  AFFINITIES   OF   THE   FAUNA.  2QI 

UNGULATA.        Bos  sondaicus. 

Tragulus  javanicus. 

Tapirus. 
EFFODIENTIA.    Manis  javanica. 

Other  forms  might  be  added  to  several  of  these  lists,  but 
those  included  are  sufficient  for  the  purpose  of  showing  that  the 
present  mammalian  fauna  of  India  is  a  complex  formed  by  an 
admixture  of  western  and  eastern  types. 

The  first  group  is  an  essentially  modern  one,  all  the  generic 
types  contained  in  it,  with  the  exception  of  Erinaceus  (which  dates 
from  the  Miocene),  being  unknown  before  the  lower  Pliocene ; 
while,  if  we  except  Erinaceus  and  Golunda,  all  occur  in  the 
Siwaliks.  In  the  Carnivora,  there  is  evidence  of  all  the  species 
except  the  first  three  being  descended  from  Siwalik  ancestors  ; 
and  it  is  quite  probable  that  the  three  species  of  Felis  may 
trace  their  origin  to  felines  which  lived  either  in  the  Siwaliks  or 
Persia  during  the  Pliocene,  in  which  event  the  lion,  and  not 
the  tiger,  should  be  regarded  as  the  characteristic  large  Indian 
feline. 

With  the  probable  exception  of  Loris,  the  second  group  is 
also  a  modern  one  ;  all  the  forms  save  Loris  and  Golunda  having 
ancestral  types  in  either  the  Pliocene  or  Plistocene  of  India,  and 
none  of  the  genera  being  known  before  the  former  epoch.  And  it 
may  be  mentioned  here  that  the  absence  of  so  many  of  the  smaller 
types  of  Oriental  mammals  from  the  Siwaliks  is  no  indication 
that  the  genera  did  not  flourish  in  India  during  the  Pliocene  age, 
but  is  due  to  the  strata  being  unsuited  to  the  preservation  of  their 
remains.  The  remarks  applicable  to  the  second  group  will  like- 
wise befit  the  third. 

On  the  other  hand,  the  fourth  and  fifth  groups  appear  to  have 
less  connection  with  the  Siwaliks,  while  several  of  the  types  are 
older  ones.  For  instance,  we  have  no  proof  of  the  existence  of 
oxen  nearly  allied  to  Bos  gaurus  in  the  Siwaliks,  although  such 
are  found  in  the  Indian  Plistocene ;  neither  is  there  any  evidence 
of  a  Siwalik  tapir.  Tupaia  is  a  near  relative  of  the  European 
Miocene  Galerix  and  the  Oligocene  Lanthanotherium ;  as  is 
Gymnura  of  the  Oligocene  Necrogymnura ;  while  Hylobates  is 

19—2 


THE   ORIENTAL   REGION.  [CHAP. 

represented  in  the  European  Miocene1.  The  reasons  for  regard- 
ing the  tiger  as  a  comparatively  modern  immigrant  into  southern 
India  have  already  been  stated.  A  Siwalik  origin  is,  however, 
indicated  for  Simla  ;  but  concerning  the  other  genera  the  palaeon- 
tological  history  is  unfortunately  a  blank. 

The  affinity  between  the  faunas  of  West  Africa  and  the  Malayan 
sub-region  has  been  already  alluded  to  ;  but  there  are  also  indica- 
tions of  a  connection  between  that  of  the  latter  area  on  the  one 
hand  and  that  of  southern  India  and  Ceylon  on  the  other,  as 
exemplified  by  the  occurrence  of  Nycticebus  in  the  Malayan  sub- 
region  and  of  Loris  in  south  India  and  Ceylon.  To  explain  the 
latter  connection,  Dr  Blanford2  has  discussed  the  possible  exist- 
ence of  a  direct  land-communication  between  the  two  areas  ;  but 
this  connection,  as  he  admits,  scarcely  seems  necessary,  since  in 
such  cases  the  true  explanation  would  seem  to  be  the  survival  of 
old  types  in  the  tropical  forest-regions.  And  here  it  may  be 
noticed  that  the  Malayan  types  common  to,  or  represented  by 
allied  forms  in  West  Africa,  are  such  as  either  have  representatives 
in  the  Indian  Pliocene  or  Plistocene,  or  such  as  we  might  naturally 
expect  to  meet  with  there  if  small  forms  were  commonly  preserved. 
For  instance,  Simia  and  Anthropopithecus,  and  Dorcatherium  and 
Tragulus,  are  all  represented  in  the  Siwaliks,  and  Atherura 
occurs  in  the  Madras  Plistocene.  This  being  so,  what  is  more 
likely  than  that  lorises,  linsangs,  and  palm-civets,  of  types  more  or 
less  intermediate  between  the  existing  Malayan  and  West  African 
representatives  of  those  groups,  should  have  flourished  in  India 
during  the  Pliocene  era  ?  Nannostiurus,  again,  should  certainly 
be  expected  to  occur  in  the  Indian  Pliocene.  On  this  point  I 
think  Dr  Wallace3  was  on  the  right  track  when,  in  writing  of  the 
Malayan  sub -region,  he  observed  that  "Here  alone,  in  the 
Oriental  region,  are  found  the  most  typical  equatorial  forms  of 
life-organisms  adapted  to  a  climate  characterised  by  uniform  but 
not  excessive  heat,  abundant  moisture,  and  no  marked  departure 
from  the  average  meteorological  state  throughout  the  year.  These 

1  If  my  memory  serves  me  right,  I  have  been  shown  teeth  from  the  upper 
Oligocene  Phosphorites  of  France  closely  resembling  those  of  Hylobates. 

2  Manual  of  Geology  of  India,  rst  Ed.  pt.  1.  p.  Ixviii. 

3  Geographical  Distribution  of  Animals,  vol.  I.  p.  335. 


VIII.]  AFFINITIES   OF   THE   FAUNA.  293 

favourable  conditions  of  life  only  occur  in  three  widely  separated 
districts  of  the  globe — the  Malay  Archipelago,  Western  Africa,  and 
equatorial  South  America.  Hence,  perhaps,  it  is  that  the  tapir  and 
trogons  of  Malacca  should  so  closely  resemble  those  of  South 
America ;  and  that  the  great  anthropoid  apes  and  crested  horn- 
bills  of  Western  Africa  should  find  their  nearest  allies  in  Borneo 
and  Sumatra." 

In  addition  to  the  resemblances  between  the  mammalian  fauna 
of  the  Malayan  sub-region  and  that  of  West  Africa,  there  are, 
however,  equally  well-marked  differences  between  the  former  and 
that  of  Ethiopia  in  general.  Among  Burmese  and  Malayan  types 
wanting  in  Africa,  we  have  especially  to  note  Tapirus,  Gymnura, 
Tupaia,  Hylobates,  and  sElurus.  From  the  small  size  of  their 
representatives  it  would  be  unfair  to  argue  anything  from  the 
absence  of  the  last  four  of  these  from  the  Siwaliks,  but  the  case  is 
very  different  with  regard  to  Tapirus,  which  ought  surely  to  have 
been  found  did  it  exist  there.  As  this  genus  is  equally  wanting 
in  the  Pikermi  and  Persian  Pliocene,  while  it  occurs  in  that  of 
France,  Germany,  England  and  China,  it  is  a  fair  inference  that  it 
has  reached  the  Malayan  countries  by  a  route  lying  north  of  India. 
And  the  occurrence  of  sElurus  in  the  English  Pliocene  suggests 
that  the  same  may  be  the  case  with  that  genus.  If  this  be  so, 
it  is  not  an  improbable  hypothesis  that  the  other  genera  men- 
tioned, all  of  which  have  representative  types  in  the  European 
Tertiaries,  may  have  migrated  eastwards  by  a  similar  route ;  and 
in  this  connection  it  is  especially  noteworthy  that  such  of  the 
genera  in  question  as  enter  India  at  all,  occur  only  in  the  eastern 
or  southern  districts. 

With  regard  to  the  date  of  the  separation  of  Ceylon  from 
India,  the  numerous  species  of  mammals  common  to  the  two 
areas  show  that  this  must  have  taken  place  at  a  very  recent  date, 
comparatively  speaking ;  although,  as  aforesaid,  at  a  period  when 
several  of  the  mammals  now  inhabiting  southern  India  had  not  yet 
occupied  that  portion  of  their  distributional  area. 

When  discussing  the  possibility  of  a  former  land-connection 
across  the  Bay  of  Bengal  between  Ceylon  and  southern  India  on 
the  one  hand,  and  the  Malayan  countries  on  the  other,  Dr  Blanford 
was  careful  to  point  out  that  the  ocean-bed  afforded  no  evidence 


294  THE   ORIENTAL   REGION.  [CHAP. 

in  favour  of  such  a  line  of  communication.  This  feature,  together 
with  certain  marked  differences  between  the  mammals  of  the  two 
areas,  appears  to  afford  a  conclusive  argument  that  these  countries 
have  never  been  much  more  closely  connected  than  they  are  at 
present.  Had  any  more  extensive  connection  existed,  we  should 
surely  expect  to  find  antelopes,  gazelles,  and  perhaps  asses,  in  the 
more  open  districts  of  upper  Burma ;  while  the  Bay  of  Bengal 
would  scarcely  have  formed  such  a  sharp  line  limiting  the  eastward 
range  of  wolves,  foxes,  hyaenas,  and  the  other  mammals  mentioned 
in  the  list  on  page  289,  as  it  actually  does.  That  list  is  confined 
to  existing  types,  but  if  fossil  forms  had  been  included,  Hippo- 
potamus might  have  been  added,  since  the  extinct  Oriental 
representatives  of  that  genus  do  not  range  further  east  than  Burma 
(whither  they  evidently  migrated  down  the  river-valleys  from 
northern  India),  no  species  being  known  from  the  Tertiaries  of 
China,  Japan,  or  the  Malayan  islands.  These  circumstances, 
together  with  the  depth  of  the  sea  in  the  Bay  of  Bengal,  seem  to 
disprove  the  suggestion  of  Dr  Wallace1  that  a  continuous  tract  of 
land  formerly  connected  Borneo  and  the  rest  of  Malaysia  with 
the  central  peak  of  Ceylon,  and  extended  eastwards  to  Hainan. 
Within  the  limits  of  the  present  volume  it  would  be  quite 
impossible  to  give  a  detailed  description  of  the 
SuWegion  mammalian  faunas  of  the  Malay  Peninsula  and 
Islands ;  and  I  have  accordingly  selected  that  of 
the  Bornean  group,  as  an  example  of  what  may  be  called  the 
typical  Malayan  sub-region,  as  distinct  from  Java,  which  differs 
markedly  in  its  fauna  from  Borneo  and  Sumatra.  The  chief 
reason  for  selecting  Borneo  is  that  its  fauna  has  been  carefully 
worked  out  by  Mr  A.  H.  Everett2  and  Mr  C.  Hose3,  from  whose 
papers  the  following  list  of  mammals  (exclusive  of  bats),  with  some 
emendations  and  additions,  has  been  compiled ;  species,  like  the 
rat,  mouse,  and  buffalo,  which  have  obviously  been  introduced, 
being  omitted.  Mr  Everett  includes  within  the  Bornean  group 
the  island  of  Palawan,  and  states  that  the  group  may  be  defined 
"  by  a  line  which  starts  from  a  point  immediately  to  the  west  of 


1  Op.  dt.  p.  359.  2  Appendix,  No.  14. 

3  Descriptive  Account  of  the  Mammals  of  Borneo,  Diss,  Norfolk,  1893. 


VIII.]  SUB-REGIONS.  295 

St  Julian  Island  in  the  Tambelan  Archipelago,  and  being  drawn 
south  of  the  Great  Natuna  (Bungoran  Island),  passes  northward  of 
Labuan  and  thence  follows  the  loo-fathom  line,  so  as  to  embrace 
Balabac,  Palawan  (Paragua),  the  Calamianes,  and  the  Cuyo 
Islands,  and,  returning  along  the  same  line  of  soundings  on  the 
southern  side  of  Palawan,  is  drawn  immediately  to  the  islands  of 
Cagayan  Sulu,  and  Sibutu, — whence  it  is  continued  through  the 
Makassar  Straits  south  of  the  Paternoster,  Lauriot  (Laset  Ketjil), 
and  Solombo  islets,  and  in  a  north-westerly  direction  through  the 
Karimata  Strait  back  to  the  island  of  St  Julian."  In  the  following 
list  the  genera  and  species  peculiar  to  the  group  are  printed  in 
italics,  those  which  are  confined  to  the  Palawan  sub-group  having 
that  name  placed  after  them  in  brackets.  The  distribution  of  the 
other  forms  has  been  indicated  as  far  as  practicable. 
PRIMATES.  Simia  satyrus.  Sumatra. 

Hylobates  leuciscus.     Java  to  Philippines. 

,,         muelleri. 

Semnopithecus  maurus.    Malay  Peninsula  and  Java. 
,,  chrysomelas. 

crudger. 
„  hosei. 

,,  everetti. 

,,  rubicundus. 

,,  frontatus. 

Nasalis  larvatus. 

Macacus  arctoides.     Burma,  China,  E.  Tibet,  etc. 
„        nemestrinus.     Burma,    Malay    Peninsula, 

Sumatra,  and  Java. 

„        cynomolgus.     Burma  to  Philippines. 
Nycticebus  tardigradus.     Burma  to  Philippines. 
Tarsius  spectrum.     Sumatra,  Java,  and  Banka. 
INSECTIVORA.    Chimarrogale  himalayica.     E.  Himalaya  and  hills 

north  of  Burma. 

Crocidura  fuliginosa.     E.  Himalaya. 
„        faitida. 
„         dor  ice. 
„         indica. 
hosei. 


296  THE   ORIENTAL   REGION.  [CHAP. 

INSECTIVORA  (cont.). 

Gymnura  rafflesi1.     S.   Tenasserim,  Malay   Penin- 
sula, and  Sumatra. 
Hylomys    suilla.     Burma,    Malay    Peninsula,    and 

Sumatra. 

Ptilocercus  lowi.    Also  in  some  neighbouring  islands. 
Tupaia  murina. 

,,       javanica.     Java  and  Malay  Peninsula. 
„        minor. 
„       gracilis. 
,,       melanura. 

,,       ferruginea.    Burma,  Malay  Peninsula,  Suma- 
tra, and  Java. 
,,       splendidula. 

„       tana.     Sumatra,  Natuna  Islands. 
,,       dor  satis. 
„      picta. 
,,       montana. 

Galeopithecus  volans.     Malay  Peninsula,  S.  Tenas- 
serim, Siam,  Sumatra,  and  Java. 

CARNIVORA.     Felis  planiceps.     Malay  Peninsula  and  Sumatra. 
,,     badia. 
„     temmincki.       E.    Himalaya,    Burma,    Malay 

Peninsula,  and  (?)  Sumatra. 
,,     bengalensis.     India  to  Philippines. 
,,     marmorata.     E.  Himalaya  to  Sumatra. 
,,     nebulosa.     E.  Himalaya  to  Formosa. 
Viverra   tangalunga.     Malay    Peninsula,    Sumatra, 

Philippines,  and  Celebes. 
Linsanga  gracilis.     Java  and  (?)  Sumatra. 
Paradoxurus   leucomystax.     Malay  Peninsula  and 

Sumatra. 
,,  hermaphroditus.     India  to  Java  and 

Sumatra. 

,,  philippinensis.     Philippines. 

Arctogale  leucotis.     Sikhim,  Burma,  Malay  Penin- 
sula, and  Sumatra. 
1  Mr  Jentink  regards  the  Bornean  form  as  a  distinct  species  (G.  alba]. 


VIII.]  SUB-REGIONS.  2Q/ 

CARNIVORA  (cent.}. 

Hemigale  hardwickei.      Malay  Peninsula  and   Su- 
matra. 
,,         hosei. 
Arctictis  binturong.     E.   Himalaya,  Burma,  Siam, 

Malay  Peninsula,  Sumatra,  and  Java. 
Helictis  everetti. 

Cynogale    bennetti.       Malay    Peninsula    and    Su- 
matra. 
Herpestes  brachyurus.     Malay  Peninsula. 

,,          semitorquatus.     Sumatra1. 
(?)  Canis  rutilans.     Malay  Peninsula,    Sumatra,    and 

Java. 
Ursus    malayanus.       Arakan,    Tenasserim,    Malay 

Peninsula,  Java,  and  Sumatra. 
Mydaus  meliceps2.     Java  and  Sumatra. 
Mustela  flavigula.     India  to  China. 

,,       nudipes.     Malay  Peninsula  and  Sumatra. 
Lutra  sumatrana.     Malay  Peninsula,  Sumatra,  and 

Java. 
,,      cinerea.     India  to  Java  and  China. 

RODENTIA.        Sciuropterus  pulverulentus.     Malay  Peninsula. 

,,  horsfieldi.     Malay  Peninsula  and  Java. 

„  setosus.     Sumatra. 

,,  genibarbis.     Java. 

,,  nigripts  (Palawan). 

Pteromys  nitidus.     Malay  Peninsula  and  islands. 

„        phceomelas. 
Rhithrosriums  macrotis. 
Sciurus    bicolor.       E.    Himalaya    to     Siam     and 

(?)  Celebes. 
„         prevosti.     Malay  Peninsula,  Sumatra,  and 

Celebes. 

,,         hippurus.     Malay  Peninsula  and  islands. 
pryeri. 

1  Jentink,  Notes  Leyden  Museum,  vol.  XVI.  p.  210  (1894). 

2  The  form  from  Calamianes  has  been  separated  as  M.  marc  Jut. 


298  THE   ORIENTAL   REGION.  [CHAP. 

RODENTIA    (cOHt.). 

Sciurus    brookei. 

„         tenuis.     Malay  Peninsula  and  islands,  to 

Siam. 

,,         lowi. 
„        jentinki. 
,,          notatus.        Malay      Peninsula,     Sumatra, 

Java,  etc. 
,,         insignis.     Malay  Peninsula,  Sumatra,  and 

Java. 

,,         ho  set. 
,,         everetti. 
,,          steerei  (Palawan). 
,,         laticaudatus.     Malay  Peninsula. 
,,         soricinus,     Java  and  Sumatra. 
Nannosciurus    exilis.      Malay    Peninsula   and   Su- 
matra. 

,,  whiteheadi, 

,,  melanotis. 

Mus  infralnteus.     Sumatra. 
,,     muelleri.     Sumatra. 
,,     sabanus. 

,,     neglectus.     Philippines. 
,,     jerdoni.     E.  Himalaya,  Tenasserim,  Java. 
,,    alticola. 

,,     ephippium.     Sumatra,  Philippines. 
,,     margaretta. 
,,     raja. 

,,     ochraceiventer. 
,,     whiteheadi, 
,,     bxodon. 
,,     baluensis. 
Chiropodomys  major. 

,,  pusillus. 

Hystrix  crassispinis . 

,,       pumila  (Palawan  sub-group). 
,,       muelleri.     Sumatra. 
Trichys  gnentheri. 


VIII.]  SUB-REGIONS.  299 

UNGULATA.       Bos   sondaicus.     Burma,    Malay   Peninsula,  Java, 

and  Bali. 

„      moellendorffi '  (Palawan  sub-group). 
Cervus  unicolor2,  var.     Probably  introduced. 
Cervulus  muntjac. 

Tragulus  napu.    S.  Tenasserim  to  Java  and  Sumatra. 
,,       nigricans  (Palawan  sub-group). 
,,       javanicus.     Malay    Peninsula    to    Cochin 

China. 

Sus  vittatus.      Java,  Sumatra,  Amboyna,  Batjian. 
,,    verrucosus.     Java,  Ceram. 
,,    barbatus*. 
„    longirostris.     Java. 
Rhinoceros  sumatrensis.     Assam  to   Siam,  Malay 

Peninsula,  and  Sumatra. 
Tapirus  indicus.     S.  Tenasserim,  Malay  Peninsula, 

and  Sumatra. 

Elephas  indicus.     Probably  introduced. 
EFFODIENTIA.  Manis  javanica  (Palawan  sub-group). 

In  this  list  the  genera  Nasalis,  Trichys,  and  Rhithrosciurus  are 
peculiar  to  the  group,  while  Ptilocercus  is  almost  so ;  and,  includ- 
ing the  latter,  there  are  no  less  than  fifty-one  species  peculiar  to 
Malaysia.  Of  these  a  very  large  number  are  common  to  Sumatra 
or  the  Malay  Peninsula,  or  both  together,  while  a  smaller  number 
occur  in  Java.  There  are  but  six  species  common  to  peninsular 
India,  among  which  the  Indian  elephant  and  the  sambar  may  have 
been  introduced  ;  but  there  are  ten  which  are  found  in  the  Eastern 
Himalaya  or  Assam.  The  most  remarkable  among  these  is  the 
Himalayan  water-shrew  (Chimarrogale  himalayica),  which  is  only 

1  Founded  on  a  skull  from  the  island  of  Busuanga,  in  the  Calamianes, 
which  probably  indicates  only  a  race  (?  introduced)  of  the  buffalo. 

2  The  so-called  C.  equinus  is  regarded  by  Dr  Blanford  as  not  specifically 
distinct   from   C.   tinicolor.     This  being  so,  it  is  probable  that  the  Bornean 
form  described  by  Mr  C.  Hose  (Ann.  Mag.  Nat.  Hist.  ser.  6,  vol.  xn.  p.  -206) 
as  C.  brookei  is  also  a  variety. 

3  Sits  ahanobarbus,  Huet,  from  Palawan,  and  S.  calamianensis,   Heude, 
from  the  Calamianes,  are  identified  by  Dr  Nehring  (Sb.  Ges.  Nattirf.  Berlin, 
1894,  pp.  190,  191)  with  this  species. 


300  THE   ORIENTAL   REGION.  [CHAP. 

found  in  the  Eastern  Himalaya,  the  mountains  north  of  Burma, 
and  Mount  Kina  Balu,  in  North  Borneo  ;  a  musk-shrew  ( Croridura 
fuliginosa\  common  to  the  Eastern  Himalaya  and  Borneo,  being 
likewise  unknown  elsewhere.  These  two  instances  alone  are 
sufficient  to  prove  that  Borneo  must  have  been  in  immediate 
connection  with  the  lands  to  the  north-west  within  the  period 
during  which  the  living  species  of  mammals  have  come  into  exist- 
ence; while  the  restriction  of  the  water-shrew  to  the  mountains 
seems  likewise  to  imply  a  former  lowering  of  the  temperature  of 
the  whole  region  sufficient  to  enable  the  creature  to  pass  from  the 
one  area  to  the  other,  or  perhaps  rather  to  have  allowed  of  its 
existence  in  the  intermediate  lowlands,  whence  it  migrated  to  its 
present  isolated  haunts.  That  Borneo  was  connected  with  the 
mainland  during  the  Pliocene  epoch  is  proved  by  the  occurrence 
in  that  island  of  the  Siwalik  Mastodon  latidens,  the  tooth  figured 
on  page  173  being  of  Bornean  origin. 

The  large  number  of  species  common  to  Borneo,  Sumatra,  and 
the  Malay  Peninsula  also  shows  that  these  three  areas  must  have 
been  very  recently  in  connection  ;  but  the  excessive  number  of 
peculiar  Bornean  forms  seems  to  indicate  that  the  former  island, 
with  the  adjacent  islets,  was  the  first  to  be  isolated.  Even  so. 
however,  the  extraordinarily  large  percentage  of  distinctive  types 
is  most  remarkable.  Regarding  the  relationship  of  the  Palawan 
sub-group  to  Borneo,  Mr  Everett  writes  that  "although  the  general 
facies  of  the  mammalian  fauna  of  the  sub-group  is  clearly  Bornean, 
it  is  to  be  noted  that  no  species  appears  to  be  peculiar  to  the 
group  as  a  whole,  a  fact  which  suggests  the  inference  that  closely 
connected  as  Borneo  has  undoubtedly  been  with  Balabac  and 
Palawan,  and  isolated  as  they  have  been  together  from  the  main- 
land of  Asia,  there  has  also  been  much  isolation  of  Borneo  and 
Palawan  inter  se" 

From  Sumatra  and  Borneo,  which  have  so  much  in  common, 
and  in  a  somewhat  less  degree  from  the  Malay  Peninsula,  Java 
differs  very  remarkably  as  regards  its  mammalian  fauna ;  a  large 
number  of  typically  Malayan  forms  being  absent,  while  others  as 
characteristically  Indian  are  present.  In  the  first  place,  the 
orangs  (Simia),  common  to  Borneo  and  Sumatra,  are  absent : 
and  the  elephant  and  tapir  are  likewise  wanting,  the  former 


VIII.]  SUB-REGIONS.  30 1 

certainly  existing  in  a  wild  state  in  Sumatra,  although  it  has  been 
considered  that  its  occurrence  in  Borneo  is  due  to  human  intro- 
duction1. Although  the  Javan  rhinoceros  (J?.  sondaicus},  as  we 
have  seen,  is  common  to  eastern  Bengal,  Burma,  and  Java,  its 
reputed  occurrence  in  either  Borneo  or  Sumatra  does  not  appear 
to  rest  upon  any  solid  basis  of  fact2;  while  the  Sumatran  species 
(R.  sumatrensis],  which  is  common  to  Borneo,  is  wanting  from 
Java.  It  has,  indeed,  been  stated3  that  certain  teeth  from  the 
Plistocene  of  Borneo  are  referable  to  the  last-named  species,  but 
the  molars  of  both  kinds  are  so  alike  that  it  is  almost,  if  not 
quite,  impossible  to  distinguish  between  them.  A  noteworthy 
circumstance  is  that  whereas  there  is  no  Siwalik  species  allied 
to  R.  sumatrensis,  yet  R.  sondaicus  is  almost  indistinguishable 
from  the  Siwalik  R.  sivalensis,  and  is  thus  proved  to  be  a  very 
ancient  Indian  type.  As  another  instance  of  the  distinctness  of 
the  mammalian  fauna  of  Java  from  that  of  Borneo  and  Sumatra, 
we  may  take  the  case  of  the  banteng  (Bos  banteng},  which  is 
wanting4  from  both  those  islands,  but  is  present  in  Java,  the 
Malay  Peninsula,  and  Burma.  Again,  the  genus  Hemigale,  of 
which  the  type  species  is  common  to  the  Malay  Peninsula, 
Sumatra,  and  Borneo,  is  quite  unknown  in  Java.  That  the  latter 
island  was  directly  connected  with  the  mainland  is,  of  course, 
proved  by  such  species  of  existing  mammals  as  are  common  to 
the  two  areas ;  but  if  further  evidence  be  needed,  it  is  to  hand  in 
the  fossil  mammals  which  have  been  obtained  from  Pati-Ajam,  in 
Java5.  These  comprise  Elephas  trigonocephalus,  E.  bombifrons, 
E.  clifti,  E.  namadicus,  E.  hysudricus,  Sus  hysudricus,  Bos  siva- 
lensis, and  Cervus  lydekkeri.  With  the  exception  of  the  first  and 
last  (which  may  prove  not  to  be  distinct  likewise),  all  these  are 
Indian  forms,  E.  namadicus  belonging  to  the  Plistocene,  while  the 
others  pertain  to  the  Siwalik  fauna;  and  it  may  be  added  that 

1  Dr  Wallace  states  that  Ursus  malayanus  is  absent  from  Java,  but  accord- 
ing to  Dr  Blanford  this  is  incorrect. 

2  See  Jentink,  Notes  Leyden  Museum,  vol.  xvi.  p.  231  (1894). 

3  Busk,  Proc.  Zool.  Soc.  1869,  p.  409. 

4  In  the  Fauna  of  British  India— Mammalia,  p.  490,  Dr  Blanford  gives 
Borneo,  and  perhaps  Sumatra,  as  part  of  the  habitat  of  the  banteng,  but  the 
animal  is  omitted  from  fauna  of  the  former  by  Mr  Everett. 

5  K.  Martin,  Sammlungen  Geol.  Reichsmusetuns  in  Leiden,  vol.  IV  (1887). 


302  THE   ORIENTAL   REGION.  [CHAP. 

the  first  three  belong  to  the  group  of  stegodont,  or  intermediate 
elephants. 

In  endeavouring  to  explain  the  relationship  of  the  Javan  fauna 
to  that  of  the  rest  of  the  Malayan  sub-region,  Dr  Wallace1  was 
first  of  opinion  that  Java,  which  was  evidently  isolated  before 
Sumatra  and  Borneo,  had  a  brief  land-connection  with  the  Siamese 
peninsula,  independently  of  those  two  islands.  This  view,  how- 
ever, was  subsequently  abandoned2,  and  the  following  hypothesis 
proposed.  From  the  evidence  of  certain  Tertiary  rocks  in  Java 
believed  to  be  of  Miocene  age,  it  is  considered  probable  that  at 
the  epoch  in  question  that  island  "would  have  been  at  least  three 
thousand  feet  lower  than  it  is  now,  and  such  a  depression  would 
probably  extend  to  considerable  parts  of  Sumatra  and  Borneo,  so 
as  to  reduce  them  all  to  a  few  small  islands.  At  some  later 
period  a  gradual  elevation  occurred,  which  ultimately  united  the 
whole  of  the  islands  with  the  continent.  This  may  have  continued 
till  the  glacial  period  of  the  northern  hemisphere,  during  the 
severest  part  of  which  a  few  Himalayan  species  of  birds  and 
mammals  may  have  been  driven  southward,  and  ranged  over  suit- 
able portions  of  the  whole  area.  Java  was  then  separated  by 
subsidence,  and  these  species  became  imprisoned  there  ;  while 
those  in  the  remaining  part  of  the  Malayan  area  again  migrated 
northward  when  the  cold  had  passed  away  from  their  former 
home,  the  equatorial  forests  of  Borneo,  Sumatra,  and  the  Malay 
Peninsula  being  more  especially  adapted  to  the  typical  Malayan 
fauna — which  is  there  developed  in  rich  profusion.  A  little  later 
the  subsidence  may  have  extended  further  north,  isolating  Borneo 
and  Sumatra,  but  probably  leaving  the  Malay  Peninsula  as  a  ridge 
between  them  as  far  as  the  islands  of  Banka  and  Billiton.  Other 
slight  changes  of  climate  followed,  when  a  further  subsidence 
separated  these  last-named  islands  from  the  Malay  Peninsula,  and 
left  them  with  two  or  three  species  which  have  since  become 
slightly  modified.  We  may  thus  explain  how  it  is  that  a  species 
is  sometimes  common  to  Sumatra  and  Borneo,  while  the  inter- 
vening island  (Banka)  possesses  a  distinct  form3." 

1  Geographical  Distribution  of  Animals,  vol.  I.  p.  359. 

2  Island  Life,  p.  360. 

3  As  exemplified  in  the  case  of  the  birds  of  the  genus  Pitta. 


VIII.]  SUB-REGIONS.  303 

Although  not  taking  into  account  the  relationship  between  the 
fauna  of  Borneo  and  that  of  the  Palawan  sub-group,  this  may 
be  accepted  as,  on  the  whole,  a  very  probable  explanation  of 
the  facts  of  the  case.  It  may  be  added  that  while  the  Javan 
rhinoceros  (£.  sondaicus]  is,  as  already  stated,  closely  allied  to  the 
Siwalik  R.  sivalensis,  the  nearest  ally  of  R.  sumatrensis  appears  to 
be  the  extinct  R.  schleiermacheri  of  the  European  Miocene,  thus 
affording  one  more  instance  of  affinity  between  the  typical  Ma- 
layan fauna  and  that  of  the  middle  Tertiaries  of  Europe.  • 

Recent  investigations  on  the  mammals  from  some  of  the  small 
chain  of  islands  lying  to  the  south-west  of  Sumatra, 
such  as  Nias  and  Sipora  in  the  Mentawi  group,  to- 
gether  with  Christmas  Island1,  lying  still  further  to     Christmas 

Islands. 

the  south,  have  shown  that  they  differ  very  markedly 
from  those   of  the  larger  islands.     From   Sipora,  in  addition  to 
bats,  Mr  O.  Thomas2  records  the  following  species,  of  which  those 
peculiar  to  the  island  are  printed  in  italics  : — 

Semnopithecus  potenziani. 

Macacus  nemestrinus.     Widely  spread. 

Tupaia  ferruginea,  var.  hypochrysa.     Java. 

Paradoxurus,  sp. 

Pteromys  nitidus.     Widely  distributed. 

Sciuropterus  lugens. 

,,          aurantiacus.     Banka. 

Sciurus  melanogaster. 
,,     fraterculus. 

Mus  siporanus. 

„     raja.     N.  Borneo. 

From  Christmas  Island  the  same  writer3  records  a  peculiar 
species  of  fruit-bat  (Pteropus  natalis],  a  variety  of  a  widely-spread 
musk-shrew  (Croddura  fuliginosa),  and  two  peculiar  rats  (Mus 
macleari,  and  M.  nativitatis],  remarkable  for  their  large  size. 
Regarding  the  bearings  of  the  faunas  of  these  islands  on  the 
general  problem  of  distribution,  Mr  Thomas  writes  that  the  collec- 

1  This    must    not  be  confounded  with  the  island  of  the  same  name  in 
Polynesia. 

2  Ann.  Mus.  Civ.  Genova,  Ser.  2,  vol.  xiv.  pp.  660 — 672  (1895). 

3  Proc.  Zool.  Soc.  1887,  pp.  511—514,  and  1888,  pp.  532 — 534. 


304  THE   ORIENTAL   REGION.  [CHAP. 

tion  from  Sipora  "does  not  show  the  very  slightest  special 
relationship  to  Sumatra,  and  therefore  lends  weight  to  the  view 
that  the  Mentawi  chain  is  the  remnant  of  a  long  peninsula  or 
island,  similar  in  shape  to,  but  separate  from  the  Malay  Peninsula 
or  Sumatra.  Further  than  this  I  cannot  at  present  go,  mainly 
because  we  know  so  little  of  the  small  terrestrial  mammals  of  the 
other  islands  of  the  chain,  those  of  the  Nicobars  being  almost 
unknown,  and  those  of  Simalu,  Sibiru,  and  Pagi  entirely,  while 
in  Nias-and  Engafio  the  collections  consist  mainly  of  bats.  Still 
the  few  indications  there  are,  such  as  the  relations  to  each  other  of 
Pteropus  nicobaricus,  modiglianii,  and  natalis,  and  of  Mus  siporamis 
and  macleari,  show  that  the  mammals,  like  the  other  animals,  pre- 
sent a  general  similarity  throughout  the  chain  the  whole  way  from 
the  Nicobars  to  Christmas  Island." 

Hitherto  the  Philippine  Islands  (exclusive  of  Palawan,  Cala- 

mianes,  etc.,  which  are  classed  with  the  Bornean 
sub-re^on.6  group ')  have  been  regarded  as  forming  a  portion  of 

the  Malayan  sub-region  ;  but  the  discovery  of  a  very 
peculiar  mammalian  fauna  in  the  mountains  of  Luzon2  clearly 
proves  their  right  to  form  a  sub-region  apart.  This  mountain 
fauna,  which  it  is  highly  probable  may  also  prove  to  be  existent  in 
Mindanao,  is  evidently  a  very  ancient  one,  showing  certain  indi- 
cations of  affinity  with  that  of  Australia;  while  the  plain  fauna 
is  of  a  more  modern  and  generally  Oriental  type.  Curiously 
enough,  the  indications  of  affinity  with  the  fauna  of  Celebes  are 
by  no  means  strongly  marked.  Unfortunately,  there  is  at  present 
no  knowledge  of  the  palseontological  history  of  the  mammalian 
fauna  of  the  group.  The  following  species  of  mammals,  exclusive 
of  bats,  have  been  recorded  from  this  sub-region3;  the  names  of 
such  genera  and  species  as  are  peculiar  being  printed  in  italic  type. 
PRIMATES.  Hylobates  leuciscus. 

Macacus  cynomolgus.     A  distinct  race4. 

Tarsius  philippinensis1 '. 

Nycticebus  tardigradus. 

1   Vide  supra,  p.  294.  2  See  Thomas,  Appendix,  No.  31. 

3  In  addition  to  the  paper  cited  above,  see  Bourns  and  Dear,  Appendix, 
No.  n.  4  Often  separated  as  M. philippinensis. 

5  Meyer,  Abh.  Mtts.  Dresden,  1894,  Art.  i,  p.  i. 


VIII.] 


SUB-REGIONS. 


305 


INSECTIVORA.    Galeopithecus  volans1. 

Tupaia  everetti. 
CARNIVORA.      Viverra  tangalunga.     Ranges  to  Celebes. 

Paradoxurus  philippinensis.     Also  Bornean. 
Felis  bengalensis. 

RODENTIA.        Sciurus  philippinensis. 
„      mindanensis. 
,,       samarensis. 
„       cagsi. 

Nannosciurus  concinnus. 
Chrotomys  whiteheadi. 
Xeromys  silaceus.          '  j 
Phlaomys  cumingi. 
,,        pallidus*. 
Crateromys  schatenbergi. 
Rhynchomys  soricoides. 
Carpomys  melanurus. 
,,        phczurus. 


Mountains  of  Luzon. 


UNGULATA. 


Batomys  granti. 
Mus  luzonicus.        \ 

,,    chrysocomus.  \ 

„    neglectus. 

,,    ephippium.     > 

Bos  mindorensis. 
Cervus  philippinus. 

,,      alfredi. 
Sus  celebensis,  var.! 


Mountains  of  Luzon. 


Mountains  of  Luzon. 


Elsewhere  only  in  Celebes. 


With  the  exception  of  the  Tarsius,  which  is  now  regarded  as  a 
peculiar  species,  the  Primates  are  all  wide-ranging  forms,  as  is  also 
the  case  with  Galeopithecus  volans,  Viverra  tangalunga,  and  Felis 
bengalensis.  A  relationship  with  Borneo  is  indicated  by  the  Para- 
doxurus and  two  species  of  Mus ;  while  the  pig  is  typically  a 
Celebean  form.  The  tamarao  (J3os  mindorensis]  has  its  nearest  ally 
in  the  anoa  of  Celebes,  but,  as  mentioned  in  an  earlier  chapter, 

\ilippinensis. 


1  The  Philippine  race  has  been  separated  as  G. 

2  Doubtfully  distinct. 

3  Equal  S.  marchei,  Huet. 


20 


306  THE   ORIENTAL   REGION.  [CHAP. 

it  is  suggested  that  the  animal  in  question  is  really  a  hybrid 
between  the  latter  and  the  Indian  buffalo.  The  two  species  of 
deer  are  small  forms  allied  to  the  race  of  the  sambar  inhabiting 
Java  and  Borneo ;  the  first  in  the  list  being  uniformly  coloured, 
while  the  second  is  spotted  with  white  at  all  ages.  The  so-called 
Cervus  marianus  of  Luzon  appears  to  be  inseparable  from  C. 
philippinus. 

It  is  remarkable  that  the  six  genera  peculiar  to  the  group  all 
belong  to  the  Muridce,  and  that  five  of  these  are  known  only  from 
the  mountains  of  Luzon.  Moreover  it  is  quite  probable  that  the 
species  from  the  latter  locality  referred  to  the  Australian  genus 
Xeromys  ought  really  to  be  generically  distinguished.  It  is  among 
the  Murtdce  that  evidences  of  Australian  affinities  are  alone  ex- 
hibited. As  these  murines  have  already  been  noticed  on  page  277 
it  will  be  unnecessary  to  allude  to  them  further  in  this  place. 

Next  to  the  peculiar  rodents  of  the  mountains,  the  most  remark- 
able feature  about  the  fauna  of  the  typical  Philippines  is  the 
absence  of  such  a  number  of  the  most  characteristic  Malayan 
genera  of  mammals.  Among  the  Primates  the  deficiency  of  orangs 
(Simia)  is  perhaps  not  very  remarkable,  but  the  total  lack  of 
langurs  (Semnopitkecus)  and  the  presence  of  only  a  single  species 
of  Macacus  and  Hylobates  are  most  noteworthy.  The  macaque  is, 
however,  distributed  over  all  the  islands  of  the  group,  and  differs 
from  other  forms  of  its  species  in  its  extremely  light  coloration,  so 
that  it  is  scarcely  likely  to  have  been  introduced  by  human  agency. 
Of  Malayan  genera  which  are  absent,  special  note  may  be  taken  of 
Linsanga,  Arctogale,  Arctictis,  Cynogale,  Herpestes,  the  wild  dogs  of 
the  sub-genus  Cyon,  Ursus,  Tragulus1,  and  Elephas.  Almost 
equally  well-marked  peculiarities  are  exhibited  by  the  bird  fauna. 

Apart  from  the  tamarao,  which  has  yet  to  be  proved  entitled 
to  rank  as  a  valid  species,  the  lowland  mammalian  fauna  of  the 
Philippines  is  such  as  might  have  well  reached  the  group  by  means 
of  a  narrow  connection  of  limited  duration  with  some  portion  of 
the  Malay  countries ;  say,  for  instance,  with  Borneo  by  way  of 
Palawan.  That  such  a  connection  must  have  been  comparatively 
recent  is  indicated  by  the  identity  of  several  of  the  Philippine 

1  Represented  by  T.  nigricans  in  Palawan. 


VIII.]  SUB-REGIONS.  307 

species  with  Malayan  forms  and  the  absence  of  any  peculiar 
genera  save  Phlceomys.  The  absence  of  such  a  number  of 
Malayan  types  indicates,  however,  either  that  the  connection  must 
have  been  exceedingly  brief,  or  that  a  large  number  of  species 
formerly  inhabiting  the  islands  have  been  destroyed  by  sub- 
mergence. On  the  other  hand,  the  presence  in  the  group  of  a 
considerable  proportion  of  widely  spread  continental  genera  of 
birds  suggests  a  more  free  communication  with  China  than  at 
present  exists ;  such  communication  having  not  improbably  taken 
place  by  way  of  Formosa.  The  mountain  fauna  of  Luzon  doubt- 
less indicates  an  earlier  type  of  colonisation. 

Note. — Since  the  foregoing  was  written  two  papers  have  appeared  on  the 
Fauna  of  the  Natuna  Islands;  viz.  O.  Thomas,  Novitates  Zool.  Vol.  n.  pp. 
26—28,  and  Thomas  and  Hartert,  I.e.  pp.  409 — 429. 


2O — 2 


CHAPTER    IX. 

THE    HOLARCTIC    REGION. 

Characteristics  of  the  Mammalian  Fauna — Mammals  of  the  Eastern  Holarctic 
Region — Plistocene  Fauna  of  the  Holarctic  Region — Geographical  Changes 
since  the  Plistocene — Western  Division  of  the  Region — Arctic  Sub -region 
— European  Sub-region — Central  Asian  Sub-region — Tibetan  Sub-region 
— Mantchurian  Sub-region  —  Mediterranean  Sub-region  —  Kashmir — Ca- 
nadian Sub-region — Transition  Zone. 

BY  far  the  most  extensive  of  all  the  zoological  regions  of  the 
globe  is  that  which  is  equivalent  to  the  whole  of  the  Palsearctic 
and  the  greater  portion  of  the  Nearctic  region  of  Messrs  Sclater 
and  Wallace,  the  one  to  which  Dr  Heilprin  (after  a  suggestion 
of  Professor  A.  Newton)  applied  the  name  Holarctic.  In  defining 
this  region,  Dr  Heilprin  separated  from  it  a  Sonoran  "transitional 
region  "  in  the  Western  Hemisphere,  and  a  similar  Mediterranean 
or  Tyrrhenian  tract  in  the  Eastern.  Of  these  the  former  is 
now  accepted  as  a  definite  region  ;  but  our  knowledge  of  the 
distribution  of  species  in  the  Eastern  Hemisphere  is  either  too 
imperfect,  or  the  interdigitation  of  the  two  faunas  is  too  complete 
to  admit  of  the  full  definition  of  a  Mediterranean  region.  Accord- 
ingly, without  prejudice  as  to  what  it  may  be  possible  to  ac- 
complish in  this  direction  in  the  future,  the  Mediterranean  area 
is  provisionally  included  in  the  Holarctic  region.  It  is,  however, 
important  to  observe  that  the  reservation  by  Dr  Heilprin  of  the 
two  transitional  tracts  already  named  justifies  the  use  of  the  term 
Holarctic  even  if  both  such  tracts  be  raised  to  the  rank  of  separate 
regions  ;  and  there  is  accordingly  no  necessity  for  the  adoption  of 
Dr  Blanford's  term  Aquilonian  as  equivalent  to  the  restricted 
Holarctic.  Used  in  the  sense  here  indicated,  the  Holarctic  region 
will  comprise  all  that  portion  of  Arctogasa  lying  north  of  the 


CHAP.  IX.]  LIMITS   AND   FEATURES.  309 


Sonoran  region  in  America,  and  of  the  Ethiopian  and  Oriental 
regions  in  the  Old  World.  The  whole  area  is  extra-tropical ;  and, 
as  Dr  Heilprin  remarks,  "  no  other  region  can  compare  with  the 
Holarctic  in  the  manifold  variety  of  its  physical  characteristics. 
Every  form  of  terrestrial  configuration,  or  condition  of  soil  and 
climate  that  may  be  represented  in  any  other  region,  is  also  repre- 
sented here,  and  on  an  imposing  scale.  From  the  ice-bound  fields 
of  the  far  north  to  the  burning  desert-wastes  of  Turkestan  on  the 
south,  and  from  the  deep  forest-grown  lowlands  to  mountain- 
summits  soaring  thousands  of  feet  above  the  level  of  perpetual 
snow,  we  pass  through  all  those  various  gradations  of  climate 
which  respectively  characterise  the  Frigid,  Temperate,  and  Torrid 
zones.  Densely  covered  forest-tracts,  supporting,  as  in  the  north, 
a  sombre  growth  of  pine  and  other  coniferous  trees,  or,  as  in  the 
south,  a  vegetation  of  almost  tropical  luxuriance,  alternate  with 
broadly  open  grass  or  pasture  lands  (tundras  of  Siberia,  American 
prairies  and  plains),  which  in  some  cases  support  over  enormous 
areas  only  a  very  scanty  vegetation,  and  in  others  display  a  profuse 
variety  of  vegetable  productions.  It  is  in  this  region  that,  in 
addition  to  a  most  bountiful  development  of  desert  tracts,  we  meet 
with  the  most  elevated  table-land  (the  Central  Asian),  and,  at  the 
same  time,  with  the  greatest  expanse  of  lowland  on  the  surface  of 
the  globe,  the  great  plain  of  Siberia  and  north-eastern  Europe." 

Although  the  essential  unity  of  the  greater  portion  of  the 
Nearctic  and  Palaearctic  regions  has  long  been  fully  recognised  by 
the  American  zoologists,  several  attempts  to  bolster  up  the  alleged 
distinctness  of  these  artificial  divisions  have  recently  been  made  in 
England1,  one  proposal  being  to  recognise  an  Arctic  or  Boreal 
circumpolar  province  cut  off  from  both  areas,  although  this  is 
practically  begging  the  whole  question. 

If  I  rightly  understand  his  view,  Dr  C.  H.  Merriam2,  who  is 
an  ardent  advocate  for  the  zoological  unity  of  the  more  northern 
portions  of  both  hemispheres,  would  distinguish  a  Boreal  circum- 
polar region  common  to  both  hemispheres  ;  while  in  America  he 
recognises  south  of  this  a  Transition  region,  followed  still  more  to 
the  south  by  the  Sonoran.  In  the  Old  World  he  would  have  an 

1  Appendix,  Nos.  28  and  35. 

2  Ibid.  No.  19,  pp.  24,  63. 


310  THE   HOLARCTIC    REGION.  [CHAP. 

analogue  of  the  Sonoran — practically  equivalent  to  the  Mediter- 
ranean sub-region  of  European  writers — but  says  nothing  about 
an  analogue  of  the  Transition;  from  which  I  infer  that  he 
would  use  the  term  Boreal  really  as  practically  equivalent  to  the 
Holarctic,  if  the  Sonoran  and  Mediterranean  areas  were  subtracted. 
In  the  New  World  the  Boreal,  exclusive  of  the  Transitional,  is 
divided  into  an  Arctic  and  a  Coniferous  Forest  Boreal  zone,  the 
latter  being  frequently  spoken  of  as  the  Boreal  "  zone,"  in  contra- 
distinction to  the  Boreal  circumpolar  "region";  the  Arctic  zone 
including  the  tract  beyond  the  limit  of  trees.  The  distinction 
between  the  Boreal  and  Transition  areas  is  certainly  not  of 
regional  value ;  and  as  the  term  Boreal  is  used  in  several  senses, 
it  had  better  give  place  to  the  earlier  Holarctic. 

As  has  been  partially  indicated  in  earlier  chapters,  and  as  will 
be  more  fully  noticed  in  the  sequel,  there  is  undoubtedly  a  marked 
distinction  between  the  mammals  of  North  America  as  a  whole 
and  those  of  Europe  and  northern  Asia,  but  this  has  been  con- 
siderably exaggerated  by  including  the  Sonoran  region  in  the  old 
Nearctic,  and  is  overshadowed  by  the  number  of  genera  and 
species  common  to  the  two  areas  and  unknown  elsewhere.  Could 
a  Mediterranean  region  be  satisfactorily  denned,  the  homogeneity 
of  the  mammalian  Holarctic  fauna  would  be  still  more  apparent ; 
but  this,  from  the  great  mingling  of  northern  and  southern  types 
which  has  taken  place  in  the  Old  World,  is,  I  think,  impracticable, 
As  has  been  already  mentioned,  it  is  probable  that  the  western 
and  eastern  halves  of  the  Holarctic  region  have  never  had  more 
than  a  comparatively  small  area  of  communication  by  way  of 
Bering  Strait,  and,  therefore,  the  further  south  we  travel  in  the 
two  areas  the  more  distinct  do  the  faunas  become;  while  only 
such  forms  as  are  capable  of  withstanding  a  certain  degree  of  cold 
have  ever  been  able  to  cross  at  all.  It  may  be  added  that  the 
evidence  for  the  unity  of  the  Holarctic  region  is  by  no  means 
solely  dependent  upon  the  mammalian  fauna,  but  is  supported  by 
many  other  groups  of  animals.  To  take  an  instance  from  the 
insects,  I  may  quote  from  Mr  W.  F.  Kirby l,  who  writes  as  follows : 
"  Had  I  been  dealing  with  Lepidoptera  only,  I  would  certainly 

1  Joitrn.  Linn.  Soc. — Zool.  1873,  p.  432. 


IX.]  CHARACTERISTICS   OF   THE   FAUNA.  311 

have  united  Dr  Sclater's  Palaearctic  region  and  Nearctic  region ; 
for  although  the  species  of  North  American  Rhopalocera  are 
seldom  identical  with  those  of  northern  Asia  and  Europe,  still  the 
genera  are  the  same  with  scarcely  an  exception,  except  a  few 
representatives  of  South  American  genera,  which  have  no  more 
right  to  be  considered  Nearctic  species  than  the  similar  chance 
representatives  of  African1  forms  in  North  Africa  or  south-west 
Europe,  or  of  Indian  forms  in  south-east  Europe,  have  to  be  con- 
sidered Palaearctic  species." 

On  the  other  hand,  North  America  differs  remarkably  from  the 
eastern  half  of  the  Holarctic  region  as  regards  its  land  molluscs. 
Thus  the  Rev.  A.  H.  Cooke2  writes,  that  "  no  district  in  the  world 
of  equal  extent  is  so  poor  in  genera,  while  those  which  occur  are 
generally  of  small  size,  with  scarcely  anything  remarkable  either  in 
colouring  or  form.  The  elongated  land-shells  (Clausilia,  Bull- 
minus]  so  characteristic  of  Europe  are  entirely  wanting,  but  a  few 
Bulimulus,  of  Neotropical  origin,  penetrate  Texas,  and  from  the 
same  sources  come  a  few  species  of  Glandina  (as  far  north  as 
South  Carolina),  Holospira  (Texas),  and  Helicina"  Probably  this 
poverty  is  largely  due  to  the  unsuitableness  of  the  greater  part  of 
North  America  to  molluscan  life ;  aided  by  the  circumstance  that 
land-molluscs  are  just  the  creatures  that  would  have  been  unable 
to  pass  over  from  Asia  by  way  of  Bering  Strait.  The  batrachians, 
again,  which  differ  most  remarkably  in  their  distribution  from 
mammals,  are  not  indicative  of  the  unity  of  the  Holarctic  region, 
the  American  types  being  very  different  from  those  of  the  Eastern 
Hemisphere. 

According,  however,  to  Mr  F.  E.  Beddard3,  "  the  earth-worms 
offer  the  best  evidence  of  any  group  in  favour  of  the  Holarctic 
region." 

Although  during  the  Plistocene  era— even  subsequently  to  the 
passing  away  of  the  glacial  period — elephants,  rhino- 
ceroses, and  hippopotami  abounded  over  the  greater     tics  ^he 
part  of  Europe,  while  species  of  the  two   former 
groups  ranged  as  far  north  as  Siberia,  and  macaques 

1  The  author  obviously  means  Ethiopian. 

2  Cambridge  Natural  History — Mollusca,  p.  339  (1895). 

3  Appendix,  No.  5,  p.  80. 


312  THE   HOLARCTIC   REGION.  [CHAP. 

were  found  in  France  and  England,  yet  the  Holarctic  region  is 
now  characterised  by  the  absence  of  all  these  animals,  save  a  few 
species  of  Macacus  on  its  southern  borders ;  and  if  a  Mediterranean 
region  could  be  satisfactorily  defined,  even  these,  as  well  as  hyaenas 
and  certain  other  southern  types,  would  likewise  be  excluded. 
The  fruit-bats  constituting  the  family  Pteropodidce.  are  likewise 
practically  wanting  in  this  region;  and  Effodientia  are  quite 
unknown.  On  the  other  hand,  carnivores,  such  as  wolves,  foxes, 
bears,  martens,  weasels,  and  the  glutton,  are  abundant ;  while  the 
rodents  are  specially  represented  by  such  types  as  the  marmots, 
beavers,  voles,  and  picas ;  and  the  ungulates  comprise  the  bisons, 
nearly  all  the  sheep,  the  true  goats  (absent  in  the  western  half  of 
the  region),  and  all  the  typical  deer. 

Referring  in  more  detail  to  the  peculiar  generic  types  common 
to  the  region  as  a  whole,  we  have,  first  of  all,  among  the  Insecti- 
vora  the  typical  or  true  shrews  (Sorex] — which  belong  to  that 
section  of  the  Soricidcz  characterised  by  having  the  tips  of  the  teeth 
stained  brownish-red — in  the  main  characteristic  of  this  region, 
although  in  America  they  extend  southwards  into  the  Sonoran. 
In  the  allied  family  of  the  Talpida,  the  mole-shrews  ( Urotrichus], 
which  are  near  relatives  of  the  European  desmans,  are  represented 
solely  by  one  Japanese,  and  a  second  North  American  species, 
although  the  latter  is  frequently  separated  generically  as  Neuro- 
trichus. 

The  peculiar  genera  of  Carnivora  are  but  few,  although  certain 
groups  are  either  confined  to,  or  very  strongly  represented  in,  the 
region.  For  instance,  among  the  Felidce  the  true  lynxes — which 
although  generally  included  in  the  genus  Felts,  are  by  some 
regarded  as  entitled  to  form  a  genus  by  themselves — are  absolutely 
confined  to  this  region,  where  they  range  as  far  south  as  Spain. 
The  bears  (Ursus],  also,  are  very  strongly  represented;  brown- 
coloured  species  being  peculiar  to  the  Holarctic  area,  as  is  the 
very  distinct  polar  species  to  its  Arctic  portions.  The  peculiar 
sea-otter,  the  sole  representative  of  the  genus  Latax,  has  a  distri- 
bution very  similar  to  that  of  the  mole-shrews ;  this  animal  occur- 
ring on  the  coasts  of  Kamschatka  and  the  Kurile  Islands,  as  well 
as  on  those  of  the  Aleutians.  The  wolverene  (Gulo},  which  is  like- 
wise the  only  member  of  its  genus,  has  a  more  extended  range, 


IX.]  RODENTS.  313 

being  found  throughout  the  forest  regions  of  northern  Europe, 
Asia,  and  North  America,  while  in  the  Plistocene  era  it  ranged  as 
far  south  as  England.  Although  marine  mammals  are  for  the 
most  part  omitted  in  this  work,  the  walruses  (Trichechidcz),  which 
now  have  a  circumpolar  distinction,  can  scarcely  be  omitted,  since 
these  animals  never  wander  far  from  land.  Remains  of  the 
existing  forms  have  been  disinterred  from  the  peat  of  the  English 
fens,  while  tusks  of  fossil  species  have  been  discovered  in  the 
Pliocene  Crag  of  the  east  of  England,  and  also  in  the  corre- 
sponding deposits  of  Belgium. 

Passing  on  to  the  rodents,  we  have  in  the  Sciuridce  the  ground- 
squirrels  or  chipmunks  ( Tamias]  practically  confined  to  the  region. 
Although  in  America  they  also  extend  into  the  Sonoran,  in  Europe 
they  are  unknown  in  the  Mediterranean  area.  The  pouches  in 
the  cheeks  for  storing  food  and  the  alternate  dark  and  light  longi- 
tudinal bands  down  the  back  serve  to  distinguish  the  chipmunks 
from  other  squirrels.  Fossil  remains  of  the  genus,  probably 
belonging  to  existing  species,  occur  in  the  Plistocene  of  Europe 
and  North  America.  The  family  of  the  beavers  (Castoridcz)  seems 
always  to  have  been  mainly  confined  to  the  Holarctic  region,  one 
of  the  two  existing  species  ( Castor  fiber]  being  European,  and 
dating  from  the  English  Plistocene,  while  the  other  (C.  canadensis] 
is  North  American.  Whereas,  however,  the  latter  ranges  south- 
wards into  the  Sonoran  region,  the  former  is  unknown  in  the 
Mediterranean  sub-region.  Fossil  species  of  this  genus  occur  in 
the  upper  Tertiaries  of  Europe  and  North  America,  where  the 
extinct  Chalicomys  is  likewise  met  with ;  but  no  beavers  are  known 
from  either  the  Siwaliks  or  the  Pikermi  beds.  Although  there  are 
few  generic  types  of  Muridce  peculiar  to  the  whole  region,  such  as 
there  are  are  important.  Foremost  of  these  are  the  great  tribe  of 
the  voles  (Microtus1},  constituting  the  typical  representatives  of  a 
sub-family  nearly  allied  to  the  cricetines,  but  distinguished  by  the 
two  longitudinal  rows  of  tubercles  on  the  crowns  of  the  molar  teeth 
being  modified  into  alternating  triangular  prisms,  and  likewise  by 
these  teeth  being  generally  of  the  hypsodont  type.  In  the  Old 
World  they  are  found  from  the  Arctic  zone  to  Asia  Minor,  while 

1  Commonly  known  by  the  later  name  of  Arvicola. 


3  H  THE    HOLARCTIC    REGION.  [CHAP. 

in  America  they  enter  the  Sonoran  region.  As  fossils,  they  appear 
to  be  first  known  from  the  upper  Pliocene  Crag  of  England ;  and 
were  thus  probably  evolved  within  the  Holarctic  region  from  the 
more  generalised  cricetines  at  a  comparatively  late  epoch.  Nearly 
allied  are  the  lemmings  (My odes),  which  are,  however,  a  more 
northern  type,  unknown  in  the  Mediterranean  sub-region,  and 
likewise  in  the  Sonoran.  Still  more  northern,  and  indeed  circum- 
polar  in  its  range,  is  the  banded  lemming,  which  alone  represents 
a  genus  (Cuniculus)  distinguished  from  the  last  by  the  absence  of 
external  ears,  the  short  and  thick  fur  on  the  feet,  the  rudimentary 
first  toe  of  the  fore  feet,  and  the  elongation  of  the  two  middle 
claws  of  the  same.  The  second  of  the  two  families  peculiar 
to  the  region  is  that  of  the  picas,  or  tailless  hares  (Lagomyidce), 
comprising  small  hare -like  animals  with  short  ears,  of  which 
all  the  living  forms  are  included  in  the  single  genus  Lagomys. 
While  the  majority  of  the  picas  are  confined  to  the  highlands 
of  Central  Asia,  some  are  found  on  the  first  snowy  range  of  the 
Himalaya,  and  both  south-east  Europe  and  the  Rocky  Mountains 
severally  possess  a  representative.  Fossil  forms  are  common  in 
the  middle  and  upper  Tertiaries  of  Europe,  as  far  south  as  Sardinia. 
Although  the  hare-tribe  (Leporidcz)  have  an  almost  cosmopolitan 
distribution,  the  majority  of  species  of  Lepus  are  inhabitants  of  the 
Holarctic  region,  Central  Asia  being  especially  rich  in  representa- 
tives of  the  genus. 

Among  the  Bovidtz,  the  bisons,  which  form  a  well-marked 
group  of  the  genus  Bos,  may  be  regarded  as  now  characteristic  of 
the  region,  although  the  American  Bos  americanus  ranges  into  the 
Sonoran.  In  addition  to  the  European  B.  bison,  which  was  for- 
merly spread  over  the  greater  part  of  Europe  and  during  the 
Plistocene  extended  into  Arctic  America,  there  is  also  the  some- 
what aberrant  yak  (B.  grunniens)  of  Tibet,  In  the  Plistocene  the 
range  of  the  group  was  somewhat  more  extensive,  remains  of  an 
extinct  species  having  been  found  in  deposits  of  that  age  in 
Texas  ;  and  there  is  likewise  another  from  the  Pliocene  of  northern 
India.  The  sheep  constitute  a  group  mainly  characteristic  of  the 
Holarctic  region ;  their  headquarters  being  the  Central  Asian 
plateau,  where  they  are  more  numerous  than  anywhere  else  in 
the  world,  although  one  species  (Ovis  vignei)  impinges  on  the 


IX.]  LIST   OF   THE    FAUNA.  315 

north-western  frontier  of  the  Oriental  region,  while  the  single 
North  American  form  also  enters  the  Sonoran.  In  a  fossil  state 
it  is  possible  that  sheep  occur  in  the  Indian  Siwaliks,  but  else- 
where they  are  first  known  from  the  Forest-bed  of  the  Norfolk 
coast,  belonging  to  the  early  part  of  the  Plistocene  epoch.  More 
nearly  allied  to  the  sheep  than  to  the  goats,  the  musk-ox  (Ovibos) 
of  Arctic  America  is  now  extinct  in  the  Old  World,  but  as  it  is 
abundant  in  the  Plistocene  of  Europe  and  Asia,  where  it  ranged  as 
far  south  as  England,  it  is  surely  entitled  to  rank  among  the  forms 
common  to  the  western  and  eastern  halves  of  the  Holarctic  region. 
Nearly  allied  extinct  species  occur  in  the  Plistocene  formations  of 
the  United  States.  Among  the  Cervidce.  there  are  three  types 
common  to  the  entire  region.  The  first,  or  Elaphine  group, 
includes  the  typical  members  of  the  genus  Cervus,  as  represented 
by  the  red  deer  (C.  elaphus)  of  the  Old  World  and  the  wapiti  (C. 
canadensis)  of  North  America;  this  group  being  characterised, 
among  other  features,  by  the  general  presence  of  both  a  brow- 
and  a  bez-tine  to  the  antlers,  although  the  latter  is  wanting  in 
the  North  African  variety  of  the  red  deer,  and  also  in  the  Tibetan 
C.  thoroldi.  The  alliance  between  the  wapiti  and  some  of  the 
forms  inhabiting  Central  Asia  is  so  close  as  to  render  it  doubtful 
whether  they  are  really  anything  more  than  varieties  of  a  single 
species.  The  single  species  of  elk  (Alces)  is  common  to  both 
halves  of  the  region ;  and  the  same  is  also  the  case  with  the 
reindeer  (jRangtfer),  which  although  now  not  found  to  the  south 
of  Europe,  ranged  during  the  Plistocene  era  into  the  south  of 
France.  The  genera  or  groups  which  may  be  regarded  as  charac- 
teristic of  the  entire  Holarctic  region  may  be  tabulated  as  follows, 
those  which  are  practically  peculiar  being  printed  in  italics : — 

Insectivora. 

SORICID^E.         Sorex.     In  America  ranges  into  Sonoran. 
TALPID^E.          Urotrichus.     Japan  and  N.  America. 

Carnivora. 

FELID^E.  Felis,  the  true  lynxes  solely  Holarctic. 

MUSTELID.E.     Latax.  \  One    species    common    to    both 

Gulo.    J         hemispheres. 
TRICHECHW&.     Trichechus. 


THE    HOLARCTIC   REGION. 


[CHAP. 


Rodentia. 


Tamtas.     Ranges  into  Sonoran. 

Arctomys. 

Spermophilus. 

CASTORID^E.     Castor.     *)    ..      0 

,,  ,,-.  \  Also  Sonoran. 

MURID^E.  Microtus. } 

Myodes. 

Cuniculus. 

LA  COM  YIDSE.    Lagomys. 

LEPORID^:.        Lepus,  the  greater  number  of  species  Hol- 
arctic. 


Ungulata. 


Bos,  the  bison  group  chiefly  Holarctic,  al- 

though   the  American   species   reaches 

the  Sonoran. 
Ovis,  just  touches  Oriental,  and  also  reaches 

Sonoran. 
Ovibos,  now  extinct  in  the  Old  World,  where 

it  was  common  in  the  Plistocene. 

The  arguments  for  the  unity  of  the  Holarctic  region  are,  how- 
ever, by  no  means  confined  to  the  case  of  genera,  for  there  are  a 
number  of  species  which  either  have  a  circumpolar  range,  or 
which  are  represented  by  closely  allied  forms  in  the  opposite 
hemisphere.  It  is  true,  indeed,  that  many  of  these  are  now  more 
or  less  exclusively  Arctic  in  their  distribution,  but  some  range  a 
long  distance  to  the  south  ;  while  during  the  Plistocene  epoch  this 
was  the  case  with  the  majority.  The  following  list  includes  the 
more  important  species  which  are  either  common  to  the  eastern 
and  western  halves  of  the  region  or  have  representative  forms 
in  the  two  hemispheres  ;  those  which  are  strictly  Arctic  having 
the  letter  P  appended  :  — 

1.  Common  lynx   (Felis   lynx).      Canadian  lynx  (F.  cana- 

densis).     P. 

2.  Wolf  (Cants  lupus). 

3.  Fox  (Cants  vulpes). 

4.  Arctic  fox  (Cants  lagopus).     P. 

5.  Brown  bear  (  Ursus  arctus). 


IX.]  LIST   OF   THE   FAUNA.  317 

6.  Polar  bear  ( Ursus  maritimus).     P. 

7.  Sea-otter  (Latax  lutris). 

8.  Pine-marten    (Mustela   martes).     American    marten   (M. 

ameHcana). 

9.  Weasel  (Mustela  vulgaris). 

10.  Wolverene  (Gulo  luscus). 

11.  Walrus  (Trichechus  rosmarus). 

12.  Arctic  vole  (Microtus  rutilus).     P. 

13.  Common  lemming  (Myodes  lemmus).    American  lemming 

(M.  obensis). 

14.  Banded  lemming  (Cuniculus  torquatus).     P. 

15.  European   beaver  (Castor  fiber}.     American  beaver  (C. 

canadensis]. 

1 6.  Mountain  hare  (Lepus  timidus^}. 

17.  Bison  (Bos  bison).     American  bison  (B.  americanus). 

1 8.  Kamschatkan  sheep  (Ovis  nivicola).     Bighorn  (O.  cana- 

densis}. 

19.  Musk-ox  (Ovibos  moschatus). 

20.  Central  Asian    deer   (Cervus  eustephanus).     Wapiti    (C. 

canadensis}. 

2 1 .  Elk  (Alces  machlis}. 

22.  Reindeer  (Rangifer  tarandus}. 

If  the  Plistocene  be  taken  into  consideration  there  may  be 
added  the  mammoth  (Elephas  primigenius)  and  the  horse  (Equus 
caballus),  remains  of  both  of  which  have  been  discovered  in 
Eschscholtz  Bay,  where  those  of  the  European  bison  also  occur. 

In  this  list  it  may  be  noticed  that  the  North  American  lynx  is 
so  closely  allied  to  the  European  form  that  it  has  been  a  question 
whether  it  is  really  more  than  a  local  variety.  Although  the 
American  wolf  has  been  separated  as  a  distinct  species,  it  is  now 
generally  identified  with  the  European  form  ;  the  same  being  also 
the  case  with  the  so-called  cross-fox  of  North  America,  which, 
together  with  another  form  from  the  Himalaya,  and  a  third  from 
North  Africa,  may  be  considered  a  mere  variety  of  the  ordinary 

1  This  name  is  commonly  applied  to  the  English  hare,  although  it  properly 
belongs  to  the  more  northern  species.  The  so-called  Polar  hare  (Z.  glacialis) 
of  Arctic  America  appears  to  be  only  a  variety. 


3l8  THE    HOLARCTIC    REGION.  [CHAP. 

fox.  Much  discussion  has  taken  place  with  regard  to  whether  any 
of  the  bears  of  North  America  are  distinct  from  the  common  brown 
bear  ( Ursus  arctus),  with  its  many  Asiatic  varieties.  According, 
however,  to  one  of  the  latest  memoirs  on  this  subject1,  all  these 
forms  appear  mere  varieties  of  the  latter,  the  so-called  cinnamon 
and  black  bears  presenting  more  distinctly  marked  differences 
than  does  the  grizzly.  The  American  marten  is  so  nearly  related 
to  the  European  marten  and  the  Asiatic  sable  that  it  is  almost 
impossible  to  point  out  valid  characters  by  which  the  three  forms 
•can  be  specifically  distinguished.  In  the  case  of  the  walrus,  the 
Pacific  form  is  distinguished  by  certain  external  features  from  the 
one  inhabiting  the  Atlantic  coasts,  although  these  are  scarcely  of 
sufficient  importance  to  be  ranked  as  specific.  In  regard  to  the 
Arctic  vole,  it  may  be  mentioned  that  although  it  is  typically  a 
polar  form,  yet  it  is  represented  in  southern  Europe  by  the  bank- 
vole  (Microtus  glareolus)  and  in  the  United  States  by  Gapper's 
vole  (M.  gapperi\  both  of  which  may  be  regarded  as  southern 
climatic  offshoots  from  the  northern  stock.  Among  the  other 
species  of  rodents  it  will  suffice  to  mention  that  the  European 
and  American  beavers  are  merely  distinguished  from  each  other 
by  the  relative  lengths  of  the  nasal  bones  of  the  skull.  And  it 
may  be  added  that  the  Kamschatkan  wild  sheep  is  so  closely 
related  to  one  race  of  the  bighorn,  or  Rocky  Mountain  sheep,  that 
it  is  very  questionable  whether  the  two  are  really  entitled  to 
specific  distinction.  The  same  is  also  the  case  with  the  two  deer 
mentioned  in  the  list.  Reference  has  already  been  made  to  the 
circumstance  that  the  musk-ox  is  extinct  in  the  eastern  division 
of  the  region.  A  few  of  the  American  forms,  such  as  the  bear, 
beaver,  bison,  and  bighorn,  enter  the  limits  of  the  Sonoran  region. 

Although,  as  will  be  shown  immediately,  there  are  a  large 
number  of  generic  types  respectively  confined  to  its  eastern  and 
western  divisions,  the  lists  given  above,  especially  the  one  relating 
to  the  species,  are  amply  sufficient  to  demonstrate  the  essential 
unity  of  the  Holarctic  region.  None  of  the  other  zoological 
regions  have  anything  like  the  number  of  common  or  representa- 

1  A.  E.  Brown,  Proc.  Ac.  Philad.  1894,  pp.  119,  129.  Merriam,  P.  BioL 
Soc.  Washington,  vol.  x.  pp.  65 — 83  (1896),  regards  the  N.  American  bears  as 
forming  several  distinct  species. 


IX.]  EASTERN    DIVISION.  319 

tive  species  which  characterise  the  two  divisions  of  the  present 
one.  It  must,  moreover,  be  remembered  that  in  the  case  of  the 
other  regions  we  have  taken  the  whole  of  the  peculiar  generic 
types  into  consideration,  although  many  of  them  are  confined  to 
small  portions  of  such  regions,  whereas  in  the  present  instance 
only  those  which  range  over  nearly  the  whole  area  have  been 
mentioned.  If,  for  instance,  we  were  to  take  the  genera  respec- 
tively confined  to  the  Indian  and  Malayan  areas  of  the  Oriental 
region,  it  would  be  found  that  the  distinction  between  the  two 
areas  would  be  nearly  or  quite  as  marked  as  are  the  two  great 
divisions  of  the  Holarctic,  while  in  the  matter  of  species  the 
differences  between  the  two  would  be  far  greater.  There  would 
accordingly  be  stronger  grounds  for  making  an  Indian  and  a 
Malayan  region  than  there  are  for  the  separation  of  a  Palsearctic 
and  a  Nearctic. 

Having  now  discussed  the  leading  mammalian  types  character- 
istic of  the  Holarctic  region  as  a  whole,  it  remains 


to   notice  those  confined   to   its  eastern   division,     t  °f 


after  which  such  as  are  restricted  to  its  western  half  Holarctic 
will  be  taken  into  consideration.  And  here  it  is 
advisable  to  repeat  that  since  the  communication  between  eastern 
Asia  and  North  America  by  way  of  Bering  Strait  appears  always 
to  have  been  of  very  limited  extent,  as  we  proceed  south  in  the 
two  great  continental  areas  the  difference  between  their  faunas 
appears  more  and  more  strongly  marked.  Accordingly,  if  it  were 
possible  definitely  to  establish  a  Mediterranean  region,  the  dis- 
tinction between  the  faunas  of  the  eastern  and  western  divisions 
of  the  Holarctic  would  be  much  less  than  is  the  case  under  the 
arrangement  here  followed. 

Passing  by  the  bats  entirely,  no  notice  will  be  taken  of  groups 
which,  like  the  hedgehogs  (Erinaceus),  are  spread  over  several  of 
the  regions  of  Eastern  Arctogaea,  since  these  are  not  in  any  way 
characteristic  of  the  eastern  division  of  the  Holarctic  region1. 
The  first  mammal  that  presents  itself  for  notice  is  accordingly  the 
water-shrew,  which  is  the  sole  representative  of  the  genus  Crosso- 
pus,  and  belongs  to  that  division  of  the  Soricidce  in  which  the 

1  Such  types  have  been  discussed  when  considering  the  fauna  of  Eastern 
Arctogaea,  supra,  p.  181. 


320  THE    HOLARCTIC   REGION.  [CHAP. 

teeth  are  stained  red  ;  the  especial  characteristic  of  the  genus  being 
the  thickly-fringed  feet  and  tail.  The  water-shrew  is  a  typical 
Holarctic  mammal,  ranging  as  far  east  as  the  Altai  mountains  and 
unknown  in  the  Mediterranean  sub-region.  In  the  second  division 
of  the  same  family,  or  that  in  which  the  teeth  are  white,  there  is  a 
peculiar  shrew  from  the  Kirghiz  steppes  nearly  allied  to  the 


FIG.   66.     RUSSIAN  DESMAN  (Myogale  moschata}. 

widely-spread  musk-shrews  (Croridura),  but  constituting  a  genus 
(Diplomesodon)  by  itself.  To  the  same  sub-family  belongs  the 
Tibetan  water-shrew  (Nectogale],  which  is  likewise  the  solitary 
representative  of  its  genus,  and  is  closely  allied  to  Chimarrogale, 
which  has,  as  elsewhere  stated,  one  Oriental  and  one  Japanese 
species.  Both  these  types  are  accordingly  closely  connected  with 
the  Oriental  region,  although  nothing  is  known  of  their  past 
history.  Far  more  characteristic  of  the  eastern  half  of  the  region 


IX.]  EASTERN    DIVISION.  321 

under  consideration  are  the  two  species  of  desman  (Myogale), 
which  are  aquatic  insectivores,  with  long  trunk-like  snouts,  some- 
what intermediate  between  the  shrews  and  the  moles.  Of  the  two 
living  species,  the  smaller  is  confined  to  the  region  of  the  Pyrenees, 
extending  as  far  north  as  the  Department  of  the  Landes,  while 
the  other  is  now  restricted  to  south-eastern  Russia,  although  its 
fossilised  remains  have  been  discovered  in  the  Plistocene  Forest- 
bed  of  the  east  of  England.  Extinct  species  occur  in  the  Mio- 
cene and  upper  Oligocene  of  the  Continent.  A  slate-coloured 
insectivore,  with  the  external  form  of  a  shrew  but  the  skull  of  a 
mole,  inhabiting  Eastern  Tibet,  constitutes  the  genus  Uropsilus ; 
and  the  more  mole-like  creature  known  as  Scaptonyx  is  likewise 
from  the  same  locality.  In  any  case,  these  two  animals  only  just 
enter  the  border  of  the  region,  so  that  they  cannot  be  regarded  as 
characteristic  Holarctic  types  ;  and,  indeed,  the  Moupin  district 
of  Eastern  Tibet  is  included  by  Dr  Wallace  in  the  Oriental  region, 
although  it  is  assigned  by  others  to  the  Holarctic.  Although  two 
species  occur  to  the  south  of  the  Himalaya,  the  true  moles  (Talpa) 
are  very  characteristic  of  the  eastern  Holarctic  region,  the  common 
species  ranging  from  England  to  Japan,  and  dating  from  the  epoch 
of  the  Norfolk  Forest-bed;  while  fossil  species,  some  of  which 
have  been  separated  as  Protalpa,  range  through  the  Tertiaries  of 
Europe  to  the  epoch  of  the  upper  Oligocene.  By  some  writers 
Talpa  moschata  of  Eastern  Tibet  is  distinguished  as  Scaptochirns. 

Among  the  Carnivora  the  widely  spread  Old  World  genera 
Genetta,  Herpestes,  and  Hyczna  enter  the  Mediterranean  sub-region, 
but  are  unknown  elsewhere  in  the  Holarctic  area  at  the  present 
day.  The  Ursidce  include  a  most  remarkable  generic  type  known 
as  ALluropus,  represented  by  the  parti-coloured  bear  of  Tibet,  in 
which  the  cheek-teeth  present  a  decided  approximation  to  the 
extinct  Hytznarctus  and  also  resemble  those  of  the  panda 
(jElurus).  This  genus  is  another  of  the  border-forms  between  the 
Holarctic  and  Oriental  regions.  On  the  other  hand,  the  badgers 
(Meles)  are  very  characteristic  of  the  area  under  consideration, 
ranging  from  England  through  the  rest  of  Europe  to  Japan  and 
China,  where  one  species  enters  the  Oriental  region,  being  found 
as  far  south  as  Hongkong.  Remains  of  extinct  badgers  occur 
in  the  lower  Pliocene  of  Persia. 

L.  21 


322  THE   HOLARCTIC   REGION.  [CHAP. 

The  list  of  more  or  less  peculiar  generic  types  among  the 
rodents  is  relatively  large.  Foremost  among  these  stands  a  very 
large  flying-squirrel  {Eupetaurus},  inhabiting  the  regions  north  of 
Kashmir  and  Tibet,  and  distinguished  from  all  other  members 
of  the  family  to  which  it  belongs  by  its  tall-crowned  (Jiypsodonf] 
cheek-teeth.  In  the  dormouse  family  (Myoxida)  the  squirrel- 
tailed  species,  which  is  the  sole  representative  of  the  genus 
~Myoxus  in  its  restricted  sense,  and  the  common  dormouse,  alone 
constituting  Muscardinus,  are  exclusively  European ;  fossil  forms 
occurring  through  the  upper  and  middle  Tertiaries. 

In  the  Muridce.  the  hamsters  (Crtcetus),  if  regarded  as  generi- 
cally  distinct  from  their  allies  the  white-footed  mice  (Sitomys]  of  the 
New  World,  are  absolutely  characteristic  of  the  eastern  division  of 
the  Holarctic  region,  where  they  range  over  a  large  portion  of 
Europe  and  northern  Asia.  Extinct  species  are  abundant  in  the 
European  Tertiaries.  Two  genera  of  the  mole-like  rodents, 
having  the  dentition  of  voles,  but  approximating  in  the  form  of 
their  body  and  limbs  to  the  moles,  constitute  a  sub-family  which  is 
also  restricted  to  this  area.  Of  these  mole-voles,  the  first  genus 
(Ellobius]  is  represented  by  one  species  from  Russia  and  another 
from  Afghanistan,  while  the  second  (Siphneus)  includes  several 
forms  from  North  and  Central  Asia;  fossil  species  of  the  latter 
having  been  described  from  the  Plistocene  of  the  Altai  and  the 
Pliocene  of  northern  China.  The  great  mole-rat  (Spalax  typhlus) 
of  southern  Europe,  Persia,  Mesopotamia,  Syria  and  Egypt,  repre- 
sents the  only  genus  among  the  Spalacidce.  which  is  confined  to 
the  present  area.  The  Dipodidce,  on  the  other  hand,  contain 
several  characteristic  eastern  Holarctic  generic  types.  The  most 
aberrant  of  these  are  the  rat-like  animals  constituting  the  genus 
Sminthus,  of  which  one  species  inhabits  eastern  and  northern 
Europe  and  Central  Asia,  while  a  second  is  found  in  Kashmir, 
and  a  third  in  the  Kansu  district  of  China.  Of  the  more  typical 
forms,  the  true  jerboas  (Dipus),  characterised  by  having  only  three 
toes,  are  exclusively  Holarctic,  ranging  from  Egypt  into  Central 
Asia,  where  they  always  frequent  desert  districts.  Of  the  genera 
with  five  toes,  Euchoretes  is  represented  by  a  single  long-snouted 
and  long-eared  species  from  the  neighbourhood  of  Yarkund  ;  and 
Platycer corny s,  which  differs  by  its  flattened  and  lancet-shaped 


IX.] 


EASTERN    DIVISION. 


323 


tail,  includes  several  species,  extending  from  Siberia  to  Nubia, 
so  that  the  genus  just  enters  the  Ethiopian  region.  It  is  repre- 
sented in  the  Plistocene  of  northern  Asia.  Lastly,  there  is  the 
genus  Alactaga,  of  which  there  are  several  species,  mostly  inhabit- 
ants of  Northern  and  Central  Asia.  The  typical  A.  jaculus 
extends,  however,  into  Persia  and  southern  Russia,  while  in  the 
Plistocene  it  ranged  as  far  west  as  Germany  ;  and  another  species 
is  an  inhabitant  of  Afghanistan.  Although  mainly  an  Ethiopian 


FIG.  67.     HEAD  OF  SPANISH  IBEX  (Capra  pyrenaica). 

and  Neotropical  family,  the  Octodontidce.  have  an  Holarctic  repre- 
sentative in  the  gundi  (Ctenodactylus\  which  inhabits  the  borders 
of  the  Sahara  in  the  neighbourhood  of  Tripoli.  The  extinct 
Pellegrinia,  of  the  Sardinian  Pliocene,  also  belongs  to  the  same 
family. 

21 — 2 


324  THE    HOLARCTIC   REGION.  [CHAP. 

Among  the  artiodactyle  ungulates  there  are  six  genera,  either 
entirely  or  mainly  confined  to  the  eastern  Holarctic  region.  Of 
these,  the  true  goats — that  is  to  say,  the  members  of  the  genus 
Capra,  as  distinct  from  the  Oriental  and  Arabian  Hemitragus — are 
almost  exclusively  Holarctic,  although  C.  walie  inhabits  the 
Abyssinian  highlands,  and  C.  siniatica,  of  Palestine  and  upper 
Egypt,  may  also  enter  the  confines  of  the  Ethiopian  region.  All 
the  goats,  it  may  be  observed,  are  essentially  mountain-dwelling 
animals,  and  the  occurrence  of  the  same  species  of  ibex  (C. 
sibiricd)  in  both  the  Altai  and  Himalaya  is  a  clear  proof  of  the 
former  prevalence  of  colder  conditions,  as  without  these  the  ani- 
mal could  not  have  passed  from  the  one  range  to  the  other.  The 
sheep  also — in  spite  of  the  existence  of  outlying  North  American 
species,  and  a  variety  of  one  Central  Asian  form  (Ovis  vignei] 
which  enters  the  north-western  confines  of  India — are  mainly  cha- 
racteristic of  the  area  under  consideration,  attaining  their  great- 
est numerical  development,  and  also  their  maximum  size,  in  the 
highlands  of  Central  Asia.  The  range  of  the  genus  includes  almost 
the  whole  of  the  eastern  Holarctic  region,  the  mouflon  (O.  musimon) 
inhabiting  the  Corsican  islands,  and  the  aberrant  arui  (O.  tragela- 
phus]  being  found  in  northern  Africa.  It  is  possible  that  fossil 
sheep  occur  in  the  Indian  Siwaliks  (where  remains  of  a  goat  allied 
to  the  Himalayan  markhor  are  also  met  with),  and  a  large  species 
is  definitely  known  from  the  Norfolk  Forest-bed.  The  next  on 
our  list  is  the  remarkable  goat-like  antelope  from  the  hills  to  the 
north  of  the  Assam  known  as  the  takin  (Budorcas],  which  is  allied 
to  the  Oriental  Nemorhczdus,  and  is  therefore  probably  an  immi- 
grant into  the  region  from  the  southward.  Allied  to  this  group  is 
the  chamois,  or  gemse  (Rupicapra),  now  confined  to  the  higher 
mountain-ranges  of  Europe  from  the  Pyrenees  to  the  Caucasus, 
but  which  during  the  Plistocene  epoch  ranged  over  many  of  the 
lowlands.  Among  the  true  antelopes,  the  addax  (Addax],  an  ally 
of  the  oryx  group,  is  an  exclusively  Mediterranean  type,  inhabiting 
North  Africa  and  Syria,  where  there  are  also  representatives  of 
other  genera  which  are  typically  Ethiopian.  More  thoroughly 
Holarctic  are  the  two  peculiar  but  allied  genera  Saiga  and  Panthol- 
ops,  each  represented  by  a  single  living  species.  The  saiga  is  now 
confined  to  the  steppes  of  western  Asia  and  Eastern  Europe,  but 


IX.] 


EASTERN    DIVISION. 


325 


during  the  Plistocene  epoch  extended  as  far  westwards  as  Germany, 
France,  and  England.  The  chiru,  as  the  representative  of  the 
second  genus  is  called,  is,  on  the  other  hand,  an  exclusively 
Tibetan  form ;  and  it  is  believed  that  a  fossil  species  occurs  in  the 
later  Tertiary  formations  of  the  same  area,  where,  curiously  enough, 
a  rhinoceros  also  existed.  Although  gazelles  (Gazella)  have  repre- 
sentatives in  both  the  Oriental  and  Ethiopian  regions,  they  are 
mainly  characteristic  of  the  desert  districts  of  the  eastern  Holarctic 
region,  being  especially  numerous  in  North  Africa,  Syria,  and  parts 


FIG.  68.     MUSK-DEER  (Moschus  moschiferus). 

of  Central  Asia.  An  inhabitant  of  the  cooler  regions  of  Asia, 
where  it  extends  from  the  south  of  Siberia  to  Kashmir  and  Cochin 
China,  the  musk-deer  (Moschus)  represents  a  peculiar  sub-family 
of  the  Cervidoe,  confined  to  the  region  under  consideration.  A 
second  species  has  been  described  from  Kansu,  in  north-western 
China,  and  it  is  not  improbable  that  the  genus  is  also  represented 
in  the  Indian  Siwaliks.  Although  agreeing  with  the  musk-deer  in 
the  absence  of  antlers  and  the  presence  of  long  tusks  in  the  upper 


326  THE    HOLARCTIC   REGION.  [CHAP. 

jaw  of  the  males,  the  Chinese  water-deer  (Hydropotes),  from  the 
valley  of  the  Yang-tsi-kiang,  belongs  to  the  more  typical  Cervidce. 
Another  genus  (Elaphodus)  more  nearly  allied  to  the  muntjacs  is 
also  Asiatic,  being  represented  by  one  species  from  near  Ningpo, 
in  China,  and  by  a  second  from  Moupin,  in  eastern  Tibet.  Of 
the  true  deer  (Cervus)  there  are  two  groups  confined  to  the  area 
under  consideration.  Firstly,  there  is  the  elaphurine  group,  repre- 
sented solely  by  the  aberrant  David's  deer  (C.  davidianus\  of 
northern  China;  and,  secondly,  we  have  the  damine  group,  of 
which  the  fallow  deer  (C.  dama)  of  the  Mediterranean  countries 
and  the  Persian  fallow  deer  (C.  mesopotamicus]  are  the  living 
forms.  Allied  types  occur  in  the  East  Anglian  Forest-bed,  and 
the  gigantic  extinct  Irish  deer  (C.  gigantens]  must  likewise  be 
included  in  the  group,  all  the  members  of  which  have  the  antlers 
more  or  less  palmated. 

As  regards  the  camels  (Camelus),  there  is  some  difficulty  in 
arriving  at  a  satisfactory  conclusion,  since  although  a  feral  race  of 
the  Asiatic  C.  bactrianus  is  met  with  in  the  deserts  bordering  Kash- 
gar,  it  is  now  pretty  well  ascertained  that  really  wild  camels  exist 
nowhere  in  the  world.  Still,  however,  as  fossil  species  of  the 
genus  are  met  with  in  the  Pliocene  of  the  Siwalik  Hills  (on  the 
borders  of  the  Oriental  and  Holarctic  regions)  and  in  the  Plisto- 
cene  of  Algeria,  it  is  probable  that  the  group  is  of  Holarctic  origin1. 

The  foregoing  survey  may  be  summarised  as  follows  :  the 
names  of  such  genera  or  groups  as  are  mainly  or  exclusively  con- 
fined to  the  eastern  division  of  the  Holarctic  region  being  printed 
in  italic  type. 

Insect!  vora. 

SORICID^:.  Crossopus. 

Diplomesodon.     C.  Asia. 
Nectogale.     Tibet. 

TALPID/E.  Uropsilus.  \    „    _.. 

_,    -V  \  E.  Tibet. 

Scaptonyx.  ) 

Talpa.     Also  enters  Oriental. 
URSID^E.  sEluropus.     Tibet. 

MUSTELID^E.       Meles.     Enters  E.  Oriental. 

1  See  page  281. 


IX.] 


EASTERN    DIVISION. 


327 


Rodentia. 


MURID/E. 


SPALA  CID&. 


OCTODONTID^E. 


Ungulata. 


CERVID^E. 


Eupetaurus.     Tibetan. 

Myoxus. 

Muscardinus. 

Cricetus. 

Ellobius. 

Sip  /incus. 

Spalax. 

Sminthus. 

Dipus. 

Euchoretes.     Central  Asian. 

Platycer  corny  s.     Enters  Ethiopia. 

Alactaga, 

Ctenodactylus.     North  African. 


CAMELID/E. 


Capra.     An  outlying  Ethiopian  species. 
Ovis.     One  N.  American  species,  and  one 

from  Central  Asia  entering  Oriental. 
Budorcas.     Tibetan. 
Rupicapra.     European. 
Addax.     Mediterranean. 
Saiga.     Central  Asian. 
Pantholops.     Tibetan. 
Gazella.    A  large  proportion  of  the  species 

E.  Holarctic. 
Cervus.      The   Elaphurine   and   Damine 

groups  exclusively  E.  Holarctic;   the 

former  Central  Asian,  and  the  latter 

Mediterranean. 

Elaphodus.     E.  Tibet  and  China. 
Hydropotes.     Chinese. 
Moschus.     Asiatic. 
(?)  Camelus. 


With  the  possible  exception  of  the  Camelidce,  none  of  the 
families  in  the  foregoing  list  are  peculiar  to  the  area  in  question— 
a  feature  presenting  a  marked  contrast  to  the  lists  of  the  character- 


328  THE    HOLARCTIC   REGION.  [CHAP. 

istic  mammalian  genera  of  the  Ethiopian  and  Oriental  regions 
given  above.  The  total  number  of  genera  which  can  in  any  sense 
be  considered  as  peculiar  to  the  eastern  half  of  the  Holarctic 
region  does  not  exceed  thirty  ;  and  among  these  Uropsilus,  Scapt- 
onyx,  Elaphodus,  and  Hydropotes  can  only  be  regarded  as  intruders 
from  the  Oriental  region  ;  while  Ctenodactylus  and  Addax  are 
manifestly  Ethiopian  types.  Indeed,  if  a  Mediterranean  region 
were  established,  the  whole  of  these,  and  probably  also  the  true 
Tibetan  forms,  would  have  to  be  removed  from  the  lists.  Apart 
from  the  absence  of  peculiar  families,  the  list  bears  no  comparison 
as  regards  numbers  with  that  of  the  mammalian  genera  distinctive 
of  the  Ethiopian  region  ;  while,  with  the  aforesaid  deductions,  it 
is  also  considerably  inferior  to  that  of  the  Oriental  region.  All 
this  confirms  the  conclusions  already  drawn  as  to  the  inad- 
visability  of  regarding  the  area  under  consideration  as  a  separate 
zoological  region. 

The  Pliocene  and  earlier  Tertiary  faunas  of  this  area  having 
already  been  considered  in  connection  with  Eastern 
Arctogsea  in  general  in  an  earlier  chapter,  we  may 


Eastern  Hoi-       pass  On  to  a  brief  review  of  the  Plistocene  mammals 

arctic  Region. 

of  the  eastern  division  of  the  Holarctic  region,  pre- 
paratory to  the  consideration  of  the  sub-regions  into  which  the 
latter  is  divided.  The  Plistocene  period  may  be  taken  in  England 
to  commence  with  the  Forest-bed  of  the  Norfolk  coast,  which 
overlies  the  topmost  of  the  Pliocene  Crag  series,  and  is  itself 
overlain  by  the  glacial  deposits.  To  a  later  epoch  of  the  same 
period  belong  the  brick-earths  and  gravels  of  our  river  valleys,  as 
well  as  the  cavern-deposits  ;  many  of  these  being  either  of  post- 
or  inter-glacial  age. 

During  the  Plistocene  period  two  very  remarkable  differences 
from  the  existing  state  of  things  have  to  be  noticed.  In  the  first 
place,  the  eastern  Holarctic  region  was  at  that  time  very  much 
less  distinctly  differentiated  from  either  the  Ethiopian  or  the 
Oriental  than  is  the  case  at  the  present  day  ;  macaques,  hyaenas, 
the  lion,  rhinoceroses,  hippopotami,  and  elephants  abounding  in 
Europe  even  as  far  north  as  England.  The  second  point  is  the 
curious  mixture  of  remains  of  existing  species  of  mammals  respec- 
tively characteristic  of  hot  and  cold  climates  met  with  in  many 


IX.]  EASTERN    PLISTOCENE   FAUNA.  329 

parts  of  England  and  France.  For  instance,  the  glutton,  rein- 
deer, Arctic  fox,  and  musk-ox  are  animals  whose  presence  seems 
indicative  of  a  more  or  less  decidedly  Arctic  climate ;  many  of  the 
voles,  picas,  jerboas,  and  susliks,  together  with  the  saiga  antelope, 
appear  to  point  with  equal  force  to  the  prevalence  of  steppe-like 
conditions  ;  while  the  hippopotamus  and  spotted  hyaena  seem  as 
strongly  in  favour  of  a  subtropical  state  of  things.  Nevertheless, 
remains  of  several  of  these  groups  have  been  found  in  such  close 
association  as  to  leave  no  doubt  that  the  animals  lived  and  died 
hard  by  where  they  are  now  buried.  Much  evidence  on  this  point 
has  been  collected  by  Sir  H.  H.  Howorth1,  who  writes  as  follows : 
"  Cuvier,  whose  prejudices  were  the  other  way,  was  long  ago  con- 
strained to  write  of  the  remains  of  reindeer  found  with  those  of 
the  mammoth  and  rhinoceros  in  the  cave  at  Breugue :  '  II  ne  faut 
pas  douter  qu'il  [the  rhinoceros]  n'ait  dte  enseveli  avec  lui  [the 
reindeer]  a  Breugue.  Ses  os  y  e'taient  pele-mele  avec  ceux  de  ce 
grand  quadrupede,  enveloppes  dans  la  meme  terre  rouge,  et 
revetus  en  partie  de  la  meme  stalactite.'  In  the  high-level 
gravels  of  the  Thames  valley  the  mammoth  and  woolly  rhinoceros 
occurred  with  the  Elephas  antiquus  ^  while  in  the  low-level  gravels 
the  Rhinoceros  leptorhinus  and  the  hippopotamus  occurred  with 
the  bison  and  the  musk-ox2,  together  with  a  worked  flint. 
The  lemming  and  the  reindeer  occurred  with  the  lion  and 
hyaena  at  Bleadon,  the  lemming  with  the  lion  and  hyaena 
at  Wookey-Hole.  The  reindeer  and  the  grizzly  bear  were 
associated  with  the  hippopotamus  at  Cefn,  and  the  E.  antiquus 
with  the  mammoth  at  Durdham  Down.  The  hippopotamus  and 
the  E.  antiquus  have  been  found  with  the  reindeer  and  bison  in 
Kirk  dale  Cave ;  the  hippopotamus  with  the  wild  boar,  the  rein- 
deer, the  mammoth,  and  the  E.  antiquus  at  Brentford.  Lartet 
says  that  in  France  remains  of  the  hippopotamus  have  been  found 
in  one  cave,  that  of  Arcy,  in  which  the  reindeer  has  also  occurred, 
accompanied  by  a  worked  flint.  At  St  Acheul  and  in  the  Somme 
valley  the  same  two  animals  have  occurred  together,  and  also  at 
Levallos,  in  the  Seine  valley.  At  Viry  Noureuil,  near  Chauny 

1  The  Mammoth  and  the  Flood  (London,  1887),  pp.  114,  115. 

2  The  author  uses  the  term  musk-sheep  for  this  animal ;  a  few  other  verbal 
alterations  have  been  made  in  the  quotation. 


33°  THE    HOLARCriC   REGION.  [CHAP. 

(Aisne),  the  mammoth  and  Rhinoceros  antiquitatis  have  occurred 
with  the  hippopotamus,  the  reindeer,  and  the  musk-ox.  At 
Bicetre,  close  to  Paris,  the  lion  is  associated  with  northern  voles, 
a  marmot,  a  lizard,  and  a  snake.  At  Montmorency  the  mole  and 
the  hedgehog  have  occurred  with  the  hamster,  the  suslik,  and  the 
pica.  In  Auvergne,  M.  Pomel  has  found  an  elephant  and  the 
woolly  rhinoceros  with  a  cat,  a  suslik,  and  a  hamster,  together  with 
snakes,  lizards,  frogs,  and  with  shells  such  as  are  still  found  in  the 
district.  In  Germany  it  is  the  same.  At  Westeregeln  the  lion 
and  the  spotted  hyaena,  the  mammoth  and  rhinoceros  were  found 
with  the  marmot,  the  suslik,  the  lemming,  the  pica,  and  the  rein- 
deer. At  Thiede,  the  mammoth,  woolly  rhinoceros,  the  horse,  the 
ox,  the  reindeer,  the  Arctic  fox,  the  lemming,  and  the  pica;  and 
so  we  might  continue  throughout  the  majority  of  the  German 
caverns." 

Many  attempts  have  been  made  to  reconcile  these  apparently 
contradictory  facts ;  one  of  the  older  views  being  that  while  the 
tropical  types  of  mammals  lived  during  warm  interludes,  they 
migrated  southwards  with  the  incoming  of  colder  conditions  to 
give  place  to  the  more  Arctic  fauna.  The  associations  mentioned 
above  render  it,  however,  perfectly  certain  that  such  an  explanation 
is  not  the  right  one.  On  the  other  hand,  it  must  be  remembered 
that  there  is  yet  much  to  be  learnt  about  the  effects  of  climate  on 
mammals  ;  and  the  mammalian  fauna  of  the  Tibetan  plateau  shews 
that  many  types  of  animals  formerly  regarded  as  more  or  less 
essentially  tropical  or  sub-tropical  are  capable  of  withstanding  a 
winter  climate  of  extreme  severity.  Thus  in  parts  of  Tibet,  as 
well  as  in  Kashmir,  langurs  and  macaques  may  be  seen  leaping 
among  the  snow-clad  branches  of  pines.  Still  it  is  perhaps  diffi- 
cult to  understand  how  two  such  animals  as  the  hippopotamus 
and  the  reindeer  could  have  inhabited  the  same  locality  contem- 
poraneously1. 

In  spite  of  this  association  of  Arctic  and  sub-tropical  forms, 
there  appears,  however,  to  be  evidence  of  a  northern  and  southern 

1  The  present  writer  is  not  prepared  to  accept  the  view  of  Mr  A.  H.  Keane 
(Mthnology,  Cambridge  Geographical  Series,  p.  65,  1896)  that  the  reindeer  has 
only  recently  become  adapted  to  a  northern  habitat.  Among  other  circum- 
stances, its  remains  are  unknown  from  the  Forest-bed. 


IX.]  EASTERN    PLISTOCENE   FAUNA.  331 

type  of  Plistocene  fauna,  England  being  apparently  somewhere 
near  the  border-line  where  the  two  met,  and,  at  times  at  least, 
overlapped  each  other.  Probably  all  or  nearly  all  of  the  living 
European  mammals  were  in  existence  at  the  same  time  ;  but  most 
of  these  need  not  be  referred  to  here,  attention  being  concentrated 
on  those  which  are  either  extinct,  or  are  now  inhabitants  of  other 
regions  or  districts.  The  following  list  includes  the  more  im- 
portant of  these  forms  ;  those  which  are  decidedly  northern  types 
being  indicated  by  a  *,  and  those  which  appear  essentially  southern 
with  a  f.  The  scientific  names  of  extinct  species  and  genera  are 
printed  in  italic,  and  of  those  still  living  in  ordinary  type. 

Primates. 

t  Barbary  Ape.  Macacus  inuus. 
f              ,,  ,,      pliocenus.     England. 

f  ,,  „       suevicus.     Switzerland. 

f  ,,  ,,       tolosanus.     France. 

Carnivora. 

fLion.     Felis  leo. 
f  Kafir  Cat.     Felis  caffra. 
t  Sabre-tooth.     Machcerodus  latidens. 
f  Spotted  Hyaena.     Hyaena  crocuta. 
t  Striped  Hyaena.  ,,       striata. 

Cave  Bear.     Ursus  spelceus. 

*  Arctic  Fox.  Canis  lagopus. 
European  Wild  Dog.         ,,      (Cyon)  europ&us. 

„        Hunting  Dog.     Lycaon  anglicus.     England. 

*  Wolverene.     Gulo  luscus. 

Rodentia. 

t  Maltese  Squirrel.     Leithia  melitensis\ 

*  Giant  Beaver.      Trogontherium  cuvieri, 

*  Northern  Vole.     Microtus  rutilus. 
Sardinian  Pica.     Lagomys  sardus. 

1  Originally  described  as  a  gigantic  dormouse,  but  shown  by  the  writer  in  a 
communication  to  the  Proceedings  of  the  Zoological  Society,  1895,  p.  860,  to  be 
allied  to  the  Sciurida. 


332  THE    HOLARCTIC   REGION.  [CHAP. 

Ungulata. 

Aurochs.     Bos  taurus,  var.  primigenius. 
*Musk-Ox.     Ovibos  moschatus. 
t  Barbary  Sheep.     Ovis  tragelaphus. 
f  Spanish  Ibex.     Capra  pyrenaica. 

English  Gazelle.     Gazella  anglica. 
t  Saiga  Antelope.     Saiga  tartarica. 

Irish  Deer.     Cervus  giganttus. 

f  Hippopotamus.  Hippopotamus  amphibius. 

fPentland's  Hippopotamus.  ,,  pentlandi. 

*  Woolly    Rhinoceros.     Rhinoceros  antiquitatis. 
Megarhine       ,,  ,,         megarhinus. 

fLeptorhine      ,,  ,,         leptorhinus. 

t  Etruscan          ,,  ,,         etruscus. 

*  Elasmothere.     Elastnotherium  sibiricum. 

*  Mammoth.  Elephas  primigenius. 
t  Straight-tusked  Elephant.       „        antiquus. 

t  Southern  Elephant.  „       meridionalis. 

(        ,,        melitensis.       \  Maltese 

T  Dwarf  Elephants.  ._  .      .  \  ,  .      , 

(        „        mnaidnensts.  I  Islands. 

t  African  Elephant.  ,,        africanus. 

In  this  list  the  Barbary  ape  is  now  confined  to  North  Africa 
and  Gibraltar.  The  lion,  although  now  restricted  to  Africa,  India, 
Persia,  and  Mesopotamia,  ranged  during  the  historic  period  into 
Thessaly ;  while  the  Kafir  cat  is  solely  African.  The  sabre- 
toothed  tiger  of  the  caves  was  the  last  survivor  of  a  genus  common 
in  the  Pliocene,  which  in  the  Plistocene  is  unknown  further  north 
than  Cromer.  The  spotted  hyaena,  whose  remains  are  so  abundant 
in  the  English  caves,  is,  as  we  have  seen  in  an  earlier  chapter,  now 
restricted  to  southern  Africa ;  while  the  striped  species,  which 
dates  from  the  upper  Pliocene,  now  ranges  from  north  Africa  to 
India.  The  cave-bear,  a  gigantic  extinct  species,  was  distinguished 
from  the  brown  bear  by  the  more  complex  structure  of  its  molar 
teeth.  Of  the  Canidce,  the  European  wild  dog  has  its  nearest  living 
ally  in  the  Altai,  the  other  species  of  the  group  being  Oriental ; 
while  the  European  hunting-dog,  which  is  known  only  by  a  single 
jaw  from  the  Glamorganshire  caves,  appears  to  have  been  closely 


IX.]  EASTERN    PLISTOCENE   FAUNA.  333 

related  to  the  living  Cape  species.  Among  the  rodents,  it  is 
only  necessary  to  mention  that  the  giant  beaver  (Trogontheriuni) 
represents  a  distinct  genus  ranging  from  the  Norfolk  Forest-bed  to 
Siberia;  and  also  that  the  Maltese  squirrel  (Leithia)  was  restricted 
to  the  islands  from  which  it  takes  its  name.  In  the  ungulates, 
the  aurochs l  was  the  gigantic  ancestor  of  the  domestic  cattle  of 
the  present  day,  but  is  unknown  living  in  a  wild  state.  The  arui, 
or  Barbary  sheep,  is  now  restricted  to  north  Africa;  while  the 
Spanish  ibex  is  confined  to  the  mountains  of  the  Iberian  penin- 
sula, its  fossil  remains  occurring  in  the  Gibraltar  caves.  The  Irish 
deer,  distinguished  by  its  great  size  and  widely-spreading  antlers, 
was  an  ally  of  the  fallow-deer,  with  which  it  is  connected  by 
means  of  another  extinct  species  or  variety  (C.  ntffi);  and  it  may 
be  mentioned  that  there  are  species  of  extinct  deer  from  the 
Forest-bed,  which  it  is  unnecessary  to  name  in  this  place.  The 
latter  deposits  are  the  source  of  the  known  remains  of  the  English 
gazelle.  The  common  hippopotamus,  which  dates  from  the  upper 
Pliocene  of  Italy,  is  now  exclusively  confined  to  Ethiopian  Africa, 
but  in  the  Plistocene  is  known  to  have  wandered  as  far  north  as 
Yorkshire.  Pentland's  hippopotamus  is  a  smaller  species  from 
Italy  and  the  Mediterranean  islands,  where  there  may  be  a  second 
still  smaller  form.  Of  the  rhinoceroses,  R.  antiqiiitatis,  which  is 
exclusively  Plistocene,  ranged  from  Central  Europe  to  Siberia ;  its 
remains  being  dredged  abundantly,  in  common  with  those  of  the 
mammoth,  from  the  Dogger  Bank,  in  the  North  Sea.  The  relation- 
ship of  this  species  to  the  extinct  Indian  Rhinoceros  platyrhinus 
and  the  living  African  R.  simus  has  been  alluded  to  in  a  previous 
chapter.  The  other  three  European  species  of  the  genus,  which, 
like  the  last,  were  two-horned  and  devoid  of  front  teeth,  date  from 
the  Pliocene,  and  form  a  group  differing  remarkably  in  dental 
characters  from  R.  antiquitatis.  While  two  of  these  species  were 
southern  types,  the  third  accompanied  the  mammoth  and  woolly 
rhinoceros  in  Siberia.  A  near  ally  of  the  rhinoceroses,  the  huge 
Siberian  Elasmotherium,  differed  remarkably  in  the  structure  of  its 
cheek-teeth,  which  are  tall-crowned,  and  shew  some  approximation 
to  those  of  the  horses. 

1  The  European  bison  is  frequently  miscalled  the  aurochs. 


334  THE    HOLARCTIC    REGION.  [CHAP. 


From  a  distributional  point  of  view,  the  European  Plistocene 
elephants  are  of  especial  interest.     Foremost  and  best-known  of 
these  is  the  mammoth  (Elephas  primigeniiis],  which,  as  stated  in 
an  earlier  chapter,  was  a  very  near  ally  of  the  existing  Indian 
species,  although  distinguished — as  we  know  from  the  evidence  of 
specimens  preserved  in  the  frozen  soil  of  Siberia — by  its  coat  of 
woolly  red  hair,  among  which  were  intermingled  long  bristly  black 
hairs.     Curiously  enough,  traces  of  this  woolly  coat  have  been 
detected  in  the  Indian  elephant,  so  that  it  is  probable  that  this 
species  originated  in  some  part  of  Asia  where  the  climate  is  colder 
than    is    that   of  India.     Regarding   the   range    of  this    species, 
Professor  Boyd  Dawkins1  remarks  that   "the  mammoth  is  very 
abundant  in  the  caverns  and  river-deposits  of  Britain  and  of  France, 
and   is   known    to  have  ranged  over  the    Pyrenees  into    Spain, 
from  the  discovery  of  specimens  in  the  zinc-mines  of  Santander. 
It  has  been  proved  by  Prof.  E.  Lartet  and  Dr  Falconer  to  have 
lived  in  the  neighbourhood  of  Rome  when  the  volcanoes  of  central 
Italy  were  active,  and  poured  currents  of  lava  and  clouds  of  ashes 
over  the  [site  of  the]  imperial  city.     It  is  common  in  northern  and 
southern  Germany,  but  it  has  not  been  found  in  Europe  north  of 
a  line  passing  through  Hamburg,  or  in  any  part  of  Scandinavia  or 
Finland.     It  occurs  in  the  auriferous  gravels  of  the  Urals  ;  and  in 
Siberia,  as  is  well  known,  it  formerly  existed  in  countless  herds, 
being  buried  in  the  morasses  in  large  numbers,  in  the  same  manner 
as  the  Irish  elks  at  the  bottom    of  the    Irish    peat-bogs.     The 
admirable  preservation  of  some  of  the  carcases  is  undoubtedly  due 
to  their  having  been  entombed    directly  after  death,  and    then 
quickly  frozen  up,  a  process  which  need  not   necessarily  imply 
climatal  conditions  unlike  those  of  the  present  time  in  Siberia." 
That   the   mammoth    ranged   across    Bering    Strait    into    Arctic 
America,  is  proved  by  the  discovery  of  its  remains  in  the  frozen 
soil  of  Eschscholtz  Bay;  but  in  the  greater  part  of  North  America 
it  was  replaced  by  the  closely-allied  E.  columbianus.     In  eastern 
Europe  there  existed  a  variety  or  species  known  as  E.  armeniaais, 
of  which  the  molar  teeth  still  more  closely  resemble  those  of  the 
Indian  elephant  than  do  those  of  the  typical  form.     The  straight- 

1  Early  Man  in  Britain  (London,  1880),  p.  106. 


IX.]  EASTERN    PLISTOCENE    FAUNA.  335 

tusked  elephant  (E.  antiquus)  is  a  more  southern  Plistocene 
type,  of  which  the  molars  are  to  a  certain  extent  intermediate 
between  those  of  the  living  Indian  and  African  species.  Still 
more  southerly  in  its  distribution  is  the  gigantic  southern  elephant 
(E.  meridionalis),  of  which  the  remains  are  found  in  the  upper 
Pliocene  of  Italy,  as  well  as  in  the  Plistocene  Forest-bed  of 
Norfolk,  and  equivalent  strata  at  Dewlish,  in  Dorsetshire.  The 
Maltese  Islands  were  the  habitat  during  the  Plistocene  epoch  of 
the  two  or  three  species  of  dwarf  elephants,  which  appear  to  have 
been  nearly  allied  to  the  existing  African  species,  but  whose  size 
was  diminished  by  the  smallness  of  the  areas  where  they  flourished. 
Lastly,  the  African  elephant,  which  is  now  restricted  to  Ethiopian 
Africa,  has  left  evidence  of  its  existence  during  the  Plistocene 
epoch  in  Algeria,  Spain,  and  Sardinia. 

The  fauna  of  the  Forest-bed  period,  among  which  the  mam- 
moth, megarhine  rhinoceros,  and  Irish  deer  are  wanting,  is,  as 
already  stated,  of  pre-glacial  age,  and,  on  the  whole,  indicative  of  a 
fairly  warm  climate,  although  there  is  some  evidence  that  the 
musk-ox  then  ranged  as  far  south  as  England.  At  the  close  of 
this  epoch,  the  southern  elephant,  together  with  a  small  bear 
known  as  Ursus  arvernensis,  appear  to  have  become  extinct.  Soon 
after,  glacial  conditions  made  their  appearance,  causing  much  dis- 
turbance and  migratory  movements  among  the  original  southern 
pre-glacial  fauna,  and  bringing  an  incursion  of  northern  forms  like 
the  reindeer,  Arctic  fox,  wolverene,  and  musk-ox,  as  well  as  of 
species  from  the  eastern  steppes  such  as  the  Saiga  antelope  and 
the  Kirghiz  jerboa  (Alactagd),  together  with  mountain  animals  like 
the  chamois,  the  ibex,  and  the  marmot,  into  the  lowlands  of  south- 
western Europe.  Among  the  northern  forms  that  then  spread 
themselves  southward  were  the  mammoth  and  the  woolly  rhino- 
ceros, which  at  this  epoch  appear  to  have  attained  their  maximum 
development. 

Unfortunately,  there  is  much  uncertainty  as  to  the  part  played 
by  the  glacial  epoch  in  the  extermination  of  the  large  mammals 
characterising  Plistocene  Europe.  By  most  English  geologists  the 
brick-earths  of  the  Thames  valley,  which  contain  remains  of  rhino- 
ceroses and  elephants  in  abundance,  as  well  as  those  of  monkeys 
more  sparingly,  are  regarded  as  of  post-glacial  age ;  but  Prof,  von 


336  THE   HOLARCTIC   REGION.  [CHAP. 

Zittel1  considers  them  pre-glacial,  or  more  probably  inter-glacial. 
If  they  are  either  inter-  or  post-glacial,  it  is  clear  that  the  cold  was 
not  the  exterminating  cause ;  and  it  is  quite  possible  that  many 
or  all  were  killed  off  by  man,  although  this  could  scarcely  be  the 
case  with  the  Siberian  fauna. 

Be  this  as  it  may,  there  is  good  evidence  that  when  northern 
forms,  such  as  the  reindeer,  wolverene,  and  banded  lemming,  had 
once  obtained  an  entrance  into  central  and  southern  Europe,  they 
remained  there  for  a  considerable  time,  since  they  were  present 
during  the  latter  portion  of  what  is  known  as  the  palaeolithic 
epoch.  With  the  advent  of  the  present  climatic  conditions  came 
in  the  present  woodland  fauna  of  central  Europe,  constituting 
what  has  been  termed  the  squirrel-  or  bison-epoch ;  and  from  that 
date,  when  animals  became  domesticated,  man  has  exercised  a 
large  influence  on  the  fauna. 

It  is  important  to  notice  that,  in  spite  of  the  mingling  of 
northern  and  southern  types  in  England,  France,  and  Germany,  to 
which  allusion  has  already  been  made,  there  seems  to  have  been 
a  distinction  between  the  northern  and  the  Mediterranean  fauna 
throughout  the  whole  of  the  later  Plistocene  epoch,  such  forms  as 
the  Barbary  sheep  and  the  fallow-deer  being  essentially  southern, 
although  the  hippopotamus,  as  we  have  seen,  extended  as  far 
north  as  Yorkshire. 

Although  the  later  Plistocene  fauna  was  spread  not  only  over 
Europe,  but  also  through  North  and  Central  Asia,  a  number  of 
the  characteristic  European  types,  such  as  the  hippopotamus,  ibex, 
chamois,  fallow-deer,  cave-bear,  and  wild  cat,  were  wanting  in 
Asia.  In  that  area  forms  are  met  with  which  are  still  character- 
istic of  the  same  districts.  As  examples  may  be  noticed  :  the 
Mongolian  gazelle  (Gazella  gutturosd],  the  Himalayan  ibex  (Capra 
sibirica),  the  Persian  wild  goat  (Capra  cegagrus),  the  argali  (Ovis 
argali),  the  musk-deer  (Moschus  moschiferus],  the  tiger  (Felts 
tigris] — of  which  the  remains  have  been  found  even  within  the 
Arctic  Circle — together  with  a  number  of  smaller  forms,  such  as 
Siphneus  aspalax,  Ellobius  talpinus,  Spalax  typhlus,  Sminthus 
vagans,  Tamias  asiaticus,  and  Mustela  zibellina.  Here,  then, 

1  Appendix,  No.  36,  p.  189. 


IX.]  LAND   CHANGES.  337 

are  clear  indications  of  a  Central  Asiatic  sub-region  as  far  back 
as  the  Plistocene  epoch. 

The  foregoing  brief  survey  of  the  Plistocene  mammals  of  the 
eastern  division  of  the  Holarctic  region  enables 
certain  deductions  to  be  drawn  as  to  geographical 
changes  which  have  taken  place  in  the  area  since  the  Plistocene- 
that  epoch. 

In  the  first  place,  the  occurrence  of  remains  of  the  tiger  in  the 
New  Siberian,  or  Liakov  Islands,  lying  far  within  the  Arctic 
Circle,  indicates  the  union  of  those  islands  with  the  Siberian  main- 
land ;  and  this  greater  extension  of  the  land  at  the  north-eastern 
extremity  of  Asia  would  naturally  lead  to  the  conclusion  that  there 
was  also  a  land-connection  with  Alaska  across  Bering  Strait. 
That  such  was  really  the  case  is  proved  by  the  discovery  during 
the  voyage  of  H.  M.  S.  "  Blossom,"  in  the  years  1825-28,  of 
remains  of  the  horse,  mammoth,  and  bison,  in  the  frozen  soil  of 
Eschscholtz  Bay,  Kotzebue  Sound,  Alaska1;  this  evidence  being 
confirmed  by  the  occurrence  of  the  musk-ox  in  the  European 
Plistocene,  as  it  is  also  by  the  number  of  species  of  mammals  still 
common  to  the  more  northern  parts  of  the  two  hemispheres.  And 
here  it  may  be  mentioned  that,  according  to  the  researches  of  the 
Russian  geologists,  Siberia,  instead  of  being  covered  like  northern 
Europe  with  a  continuous  ice -sheet  during  the  glacial  epoch, 
had  only  a  number  of  comparatively  small  glaciers,  so  that  the  pre- 
glacial  fauna  was  able  to  exist  here  at  a  time  that  it  could  not  live 
in  Europe.  Still  the  frozen  condition  of  the  subsoil,  and  the 
formation  of  ground-ice  in  the  rivers,  rendered  the  preservation  of 
the  carcases  of  mammoths,  rhinoceroses,  bison,  and  musk-oxen  an 
easy  matter. 

Passing  to  south-western  Europe,  the  occurrence  of  remains  of 
the  African  elephant  in  Sicily  and  Spain,  together  with  the 
presence  of  small  allied  species  in  the  Plistocene  of  Malta,  and 
likewise  of  remains  of  the  Barbary  sheep  and  Barbary  ape  in 
southern  Europe,  indicates  a  free  land-communication  between 
Europe  and  Africa,  both  by  way  of  the  Straits  of  Gibraltar,  and 
likewise  between  Italy,  Sicily,  and  Tunis;  Malta  being  then  also  in 

1  Beechey's  Voyage  to  the  Pacific  and  Behring's  Straits  in  H.M.S. 
"Blossom,"  Vol.  II. 

L.  22 


THE   HOLARCTIC   REGION.  [CHAP. 

connection  with  the  mainland.  Probably  also  it  was  by  one  or 
both  of  these  routes  that  the  hippopotamus  and  spotted  hyaena 
passed  between  Europe  and  Africa,  as  it  is  scarcely  likely  that  the 
former  animal,  at  least,  travelled  round  by  way  of  Turkey  and 
Syria.  Writing  of  the  Maltese  islands,  Leith-Adams1  observes 
that  "although  from  their  smallness  the  islands  furnish  only  scant 
evidences  of  the  complicated  and  extensive  oscillations  of  level 
to  which  the  original  area  has  been  subjected  from  first  to  last, 
nevertheless  the  data  I  have  furnished  are  at  the  least  suggestive, 
and,  in  conjunction  with  the  fossilised  remains,  seem  to  lead  to 
the  belief,  that  in  the  first  place  there  was  an  upheaval  of  a  large 
tract  of  land  in  this  portion  of  the  Mediterranean  at  some  period 
towards  or  after  the  close  of  the  Miocene  epoch.  In  the  second 
place,  that  during  the  Quaternary  [Plistocene]  period,  the  whole, 
or  at  least  all  excepting  perhaps  the  tops  of  the  Benjemma  heights 
and  Gozo  hills,  were  again  submerged  ;  and,  thirdly,  that  a  re- 
elevation  of  the  land  took  place,  ending  in  the  present  insular 
fragments.  Perhaps  in  the  first  case  there  was  a  connection  or 
contemporaneity  in  the  upheaval  of  the  Miocene  beds  of  Malta, 
Sicily,  Italy,  Candia,  the  Red  Sea,  Egypt,  Arabia,  Cerigo,  Azores, 
Algeria,  Southern  France,  and  Spain.  Thus  the  islands  of  the 
inland  sea  may  represent  portions  of  a  land  area  now  occupied 
more  or  less  by  water.  When  this  area  began  to  sink  is  not 
apparent,  but  the  fact  that  the  same  elephant  and  hyaena  now 
living  in  Africa  existed  in  Sicily,  shews  that  there  was  a  land- 
connection  between  the  two  at  a  very  recent  epoch." 

Regarding  the  nature  of  the  former  connection  between  Italy, 
Sicily,  and  Malta,  Dr  Wallace2  writes  that  a  comparatively  shallow 
sea  or  submerged  bank  incloses  Malta  and  Sicily,  and  "  that  on 
the  opposite  coast  a  similar  bank  stretches  out  from  the  coast  of 
Tripoli,  leaving  a  narrow  channel,  the  greatest  depth  of  which  is 
240  fathoms.  Here,  therefore,  is  a  broad  plateau,  which  an  eleva- 
tion of  about  1,500  feet  would  convert  into  a  wide  extent  of  land 
connecting  Italy  with  Africa ;  while  the  same  elevation  would  also 
connect  Morocco  with  Spain,  leaving  two  extensive  lakes  to  repre- 


1  The  Nile  Valley  and  Malta  (London,  1870),  p.  211. 

2  Geographical  Distribution  of  Animals,  Vol.  I.  p.  201. 


IX.]  LAND   CHANGES.  339 

sent  what  is  now  the  Mediterranean  Sea,  and  affording  free  com- 
munication for  land  animals  between  Europe  and  North  Africa." 

Probably  the  dwarf  elephants  of  Malta  were  developed  from  a 
larger  form,  closely  allied  to  or  identical  with  the  African  elephant, 
after  the  separation  of  the  island  itself  from  the  mainland.  With 
regard  to  Leith-Adams'  idea  of  the  subsequent  submergence  of 
Malta,  it  is  pretty  certain  that  this  could  not  have  been  complete, 
since  that  island  is  inhabited  by  a  large  species  of  weasel  (Mustela 
africand)  common  to  Egypt,  and  perhaps  the  south  of  Italy1; 
this  animal  being  doubtless  a  survivor  from  the  old  fauna  of  the 
Plistocene  land  connecting  Italy,  Sicily  and  Malta  with  northern 
Africa. 

In  north-western  Europe  there  are  equally  conclusive  evidences 
of  the  connection  of  the  British  Islands  with  the  Continent  during 
the  Plistocene  epoch.  On  many  parts  of  the  English  coasts  there 
occur,  for  instance,  submerged  forests  dating  from  a  comparatively 
recent  epoch,  which,  when  exposed  during  exceptionally  low  tides, 
are  seen  to  contain  the  stumps  of  trees  in  their  original  upright 
position,. and  with  their  roots  still  implanted  in  the  soil.  Forests 
of  this  kind  are  found  near  Torquay  and  Falmouth,  as  well  as  on 
several  parts  of  the  Welsh  coasts  and  in  Holyhead  harbour ;  the 
submergence  which  has  taken  place  in  the  case  of  the  one  at 
Falmouth  being  estimated  at  upwards  of  70  feet.  Again,  the  pre- 
glacial  Norfolk  Forest-bed,  so  often  alluded  to  in  the  foregoing 
pages,  affords  evidence  of  an  extensive  submergence  on  the  east 
coast  of  England ;  this  being  supplemented  by  the  Dogger  Bank 
in  the  North  Sea,  from  which,  as  already  mentioned,  such  numbers 
of  remains  of  the  mammoth,  as  well  as  those  of  the  woolly  rhino- 
ceros and  other  mammals,  have  been  dredged.  Additional  evidence 
in  favour  of  the  same  subsidence  is  afforded  by  the  numerous 
ancient  river-channels  and  valleys  found  in  many  parts  of  Britain, 
which  are  situated  at  depths  of  from  one  to  two  hundred  feet 
below  the  present  level  of  the  land,  and  frequently  cut  right  across 
the  existing  drainage  lines,  so  as  to  connect  valleys  now  com- 
pletely distinct.  These  ancient  channels,  which  are  now  completely 
choked  with  sand,  mud,  or  gravel,  have  only  been  revealed  by  the 

1  See  Thomas,  Proc.  Zool.  Soc.  1895,  pp.  128 — 131. 

22—2 


340  THE    HOLARCTIC   REGION.  [CHAP. 

aid  of  the  borer,  but  their  evidence  is,  nevertheless,  unimpeach- 
able. 

From  these  and  other  lines  of  evidence,  we  learn  that  during 
the  Plistocene  epoch  not  only  was  England  connected  with  France 
across  the  English  Channel,  but  that  the  land  extended  up  the 
North  Sea  at  least  as  far  as  the  Dogger  Bank ;  the  Ouse,  the 
Thames,  the  Rhine,  and  perhaps  the  Elbe  originally  uniting  to 
form  one  mighty  river,  discharging  far  up  in  the  North  Sea. 
During  a  portion  of  this  period  Ireland  was  in  connection  with 
the  British  Islands;  and  it  has  been  suggested  by  Leith- Adams1 
that  the  connection  was  with  Scotland,  owing  to  the  circumstance 
that,  with  the  exception  of  the  cave-bear,  all  the  living  and  extinct 
Irish  mammals  have  been  recorded  from  Scotland,  while  a  number 
of  the  English  Plistocene  mammals  appear  never  to  have  reached 
the  latter  country.  On  the  other  hand,  Dr  R.  F.  Scharff 2,  from  a 
study  of  the  freshwater  fishes  and  molluscs,  is  of  opinion  that 
"  Ireland  was  in  later  Tertiary  times  connected  with  Wales  in  the 
south  and  Scotland  in  the  north  ;  whilst  a  freshwater  lake  occupied 
the  present  central  area  of  the  Irish  Sea.  The  southern  connec- 
tion broke  down  at  the  beginning  of  the  Plistocene  period,  the 
northern  connection  following  soon  after.  There  is  no  evidence 
of  any  subsequent  land-connection  between  Great  Britain  and 
Ireland."  Since  the  above  was  written  Dr  Scharff  (Mem.  Soc. 
ZooL  France,  vol.  vin.  pp.  436-474,  1895)  has  further  developed 
his  views  on  the  origin  of  the  Irish  fauna.  He  concludes  that 
all  the  Irish  mammals  reached  the  island  in  the  early  Plistocene 
(Forest-bed);  such  British  forms  as  are  unknown  in  Ireland  being 
considered  to  have  reached  Britain  later,  when  Ireland  was 
isolated. 

Further  reference  to  the  former  connection  or  connections 
between  Britain  and  the  Continent  will  come  more  conveniently 
later. 

The  generic  and  specific  mammalian  types  common  to  the 
eastern  and  western  divisions  of  the  Holarctic  region 

Western 

Division  of  the     having  been  already  referred  to,  we  may  at  once 
proceed  to  the  consideration  of  those  characteristic 

1  Proc.  Roy.  Irish  Acad.  Ser.  2,  Vol.  III.  (1883). 

2  Appendix,  No.  25. 


IX.]  WESTERN   DIVISION.  34! 

of  the  western  division.  And  here  it  may  be  mentioned  in  respect 
to  the  two  areas,  that  whereas  many  generic  types  of  animals  were 
unable  to  pass  from  the  one  to  the  other  owing  to  the  high 
latitude  of  the  strip  of  connecting  land,  yet  in  other  cases  the 
geographical  limits  of  the  range  of  certain  genera  in  the  Old  World 
form  also  an  important  factor  in  the  case.  As  stated  a  few  para- 
graphs back,  many  of  the  characteristic  European  Plistocene 
mammals,  such  as  the  hippopotamus,  the  fallow-deer,  and  the 
cave-hyaena,  never  extended  into  the  Asiatic  portion  of  the  Hoi- 
arctic  region,  so  that  these  and  many  other  forms  never  could 
have  had  an  opportunity  of  crossing  Bering  Strait,  even  had 
they  been  capable  of  existing  in  such  a  high  latitude. 

Excluding  bats  and  seals,  the  following  genera  of  mammals 
will  be  found  confined  to  the  western  half  of  the  Holarctic  region, 
although  some  of  these  range  southwards  into  the  Sonoran.  There 
are  also  certain  genera  which  appear  to  be  typically  Sonoran, 
whose  range  includes  part  of  the  western  Holarctic  region,  but 
these  are  best  considered  in  the  light  of  intruders  from  the 
south l. 

Among  the  shrews  of  the  western  Holarctic  there  are  two 
species,  viz.  Sorex  palustris,  of  the  Rocky  Mountains,  and*S.  hydro- 
dromus,  of  Unalaska  Island,  which  differ  from  all  their  allies  in  the 
presence  of  long  fringes  of  hair  to  the  feet,  although  they  resemble 
ordinary  species  of  the  genus  in  the  characters  of  their  dentition 
and  tail.  In  consequence  of  these  differences  these  aquatic  shrews 
have  been  referred  by  some  writers  to  a  separate  genus,  under  the 
name  of  Neosorex ;  although  such  distinction  is  considered  by  Dr 
Merriam  unnecessary.  There  is,  however,  one  genus  of  Insectivora 
(Condylura),  represented  by  the  star-nosed  mole,  absolutely  charac- 
teristic of  this  area.  Allied  in  structure  and  habits  to  the  Old 
World  moles,  which  are  totally  wanting  in  America,  this  animal 
takes  its  name  from  the  presence  of  a  star-like  ring  of  fleshy 
appendages  at  the  extremity  of  the  muzzle. 

The  Carnivora  include  no  peculiar  genera2;  but  the  Rodentia, 

1  It  may  be  well  to  mention  here  that  the  majority  of  American  zoologists 
regard  as  genera  a  number  of  groups  to  which  the  present  writer  would  not  be 
disposed  to  grant  more  than  sub-generic  rank. 

2  Mephitis,  Taxidea,  etc.,  appear  to  be  of  Sonoran  origin. 


342  THE    HOLARCTIC   REGION.  [CHAP. 

which,  as  in  the  eastern  division,  are  very  numerous,  comprise  one 
family,  as  well  as  several  genera,  restricted  to  this  area.  In  the 
Sciuridce  the  marmot- like  genus  Cynomys  ranges  into  the  Hoi- 
arctic,  but  is  considered  by  Dr  Merriam  as  chiefly  characteristic  of 
the  Sonoran  region;  and  the  same  is  the  case  with  the  white- 
footed  mice  (Sitomys) — of  which  there  is  but  a  single  Holarctic 
representative,  while  the  Sonoran  species  are  very  numerous — and 
also  with  the  wood-rats  (Neotoma),  of  which  a  sub-genus  is 
restricted  to  the  Holarctic.  The  family  peculiar  to  the  region  is 
that  of  the  Haplodontidtz,  or  sewellels,  represented  by  two  species 
of  the  genus  Haplodon,  from  the  districts  west  of  the  Rocky 
Mountains.  Closely  allied  to  the  squirrels,  these  rodents  are 
distinguished  from  the  latter  by  the  absence  of  postorbital  pro- 
cesses to  the  frontal  bones  of  the  skull,  the  depressed  skull,  and 
the  rootless,  or  hypsodont,  cheek-teeth ;  all  these  characters  indi- 
cating a  more  specialised  type.  In  the  Muridce,  the  voles  of  the 
genus  Phenacomys  connect  the  more  typical  members  of  the 
group  with  cricetines  like  the  wood-rats  (Neotoma).  Several 
species  have  been  described.  A  more  southern  type  is  the  single 
representative  of  the  allied  genus  Synaptomys,  distinguished  by  its 
grooved  upper  incisors  \  its  molar  teeth  resembling  those  of  the 
lemmings,  while  its  skull  is  of  the  same  structure  as  in  the  true 
voles.  According  to  Dr  Merriam,  this  animal  is  restricted  to  the 
southern  part  of  the  Holarctic  area,  or  what  he  terms  the  Transi- 
tion region.  In  the  same  great  family  the  well-known  aquatic 
musk-rat,  or  musquash  (Fiber),  may  be  considered  an  Holarctic 
type,  since  it  is  found  in  the  "  barren-grounds  "  on  the  borders  of 
the  Arctic  sea,  although  it  ranges  southwards  into  the  Sonoran. 
Closely  allied  to  the  voles,  with  which  it  agrees  in  the  characters 
of  the  skull  and  teeth,  this  animal  differs  by  the  long,  compressed, 
nearly  naked,  and  reticulate  tail ;  the  naked-soled  feet  being  partly 
webbed,  and  the  whole  body  adapted  to  an  aquatic  mode  of  life. 
Its  fossil  remains  occur  in  the  Plistocene  of  the  United  States.  It 
is  noteworthy,  as  a  negative  characteristic  of  the  Holarctic  area, 
that  no  members  of  the  exclusively  New  World  family  Geomyidce 
are  found  within  its  limits.  On  the  other  hand,  in  the  family 
Dipodida,  the  j  urn  ping-mice  of  the  genus  Zapus,  of  which  several 
species  are  recognised  by  North  American  zoologists,  are  solely 


IX.] 


WESTERN    DIVISION. 


343 


Holarctic,  the  typical  Z.  hudsonianus  dating  from  the  Plistocene 
epoch.  A  distinctive  feature  of  the  western  Holarctic  region  is 
the  absence  of  true  porcupines  (Hystrix),  their  place  being  taken 
by  the  Canadian  porcupine  (Erethizon],  which  belongs  to  the  same 
sub-family  as  the  South  American  porcupines,  although  distin- 
guished, among  other  characters,  by  its  short  and  non-prehensile 
tail.  It  is  a  native  of  the  wooded  portions  of  Canada  and  the 
United  States,  and  its  remains  have  been  discovered  in  a  cave  in 
Pennsylvania. 


FIG.  69.     ROCKY  MOUNTAIN  GOAT  (Haploceros  montamis] 

Among  the  ungulates,  the  remarkable  animal  known  as  the 
Rocky  Mountain  goat,  which  alone  represents  the  genus  Haplo- 
ceros, and  differs  from  all  other  ruminants  by  the  extreme  shortness 
of  the  cannon-bone  in  both  the  front  and  hind  limbs,  is  exclusively 
an  inhabitant  of  the  western  Holarctic  region.  The  same  is  now 


344  THE   HOLARCTIC   REGION.  [CHAP. 

the  case  with  the  musk-ox  (Ovibos),  but  as  this  animal  ranged  over 
Europe  and  northern  Asia  during  the  Plistocene,  it  can  scarcely  be 
regarded  as  distinctive  of  the  western  area.  Of  other  peculiar 
New  World  ungulates,  the  prong-buck  (Antilocapra)  and  certain 
deer  of  the  genus  Cariacus  are  found  within  the  Holarctic  region, 
but  the  former  seems  to  be  essentially  a  Sonoran  type,  while  the 
latter,  although  probably  also  of  Sonoran  origin,  occurs  through 
Central  and  South  America. 

The  Tertiary  genera  of  mammals  peculiar  to  North  America 
may  be  best  considered  in  the  chapter  devoted  to  the  Sonoran 
region,  to  which  they  for  the  most  part  belong ;  and  this  portion 
of  the  subject  may  be  accordingly  concluded  by  tabulating  the 
existing  genera  or  groups  peculiar  to  the  area  under  consideration. 
These  will  stand  as  follows,  viz. : — 

Insectivora. 

SORICID.E.     Sorex.     The  sub-genus  or  genus  Neosorex. 
TALPID^E.      Condylura. 

Rodentia. 

HAPLODONTIDJE.     Haplodon. 
MURID^:.  Phenacomys. 

Synaptomys.     Confined  to  the  southern 
portion  of  the  area. 

Fiber.     Enters  Sonoran. 
DIPODID^E.  Zapus. 

HYSTRICID.E.  Erethizon. 

Ungulata. 

BOVID^E.  Haploceros. 

Even  if  we  add  to  the  above  certain  other  sub-generic  types, 
such  as  the  spruce-squirrels  (Tamiasdurus)  and  the  bushy-tailed 
wood-rats  (Teonoma),  and  likewise  take  into  account  the  number 
of  Old  World  types  (in  many  cases  widely-distributed  ones)  that 
are  absent,  it  can  scarcely  be  urged  that  such  an  assemblage  is 
sufficient  to  constitute  a  zoological  region  by  itself.  Those  of  my 
readers  desirous  of  consulting  lists  of  the  species  inhabiting  the 
Arctic  and  Boreal  zones  of  Dr  Merriam,  will  find  them  in  his 
memoir1. 

1  Appendix,  No.  19,  pp.  24,  25. 


IX.]  WESTERN   DIVISION.  345 

In  America,  probably  owing  to  the  north  and  south  trend  of 
the  mountain-ranges,  the  glacial  period  has  had  an  even  more 
marked  effect  than  in  the  Old  World.  On  this  subject  Dr 
Merriam1  writes  that  "not  only  are  the  pre-Plistocene  animals  and 
plants  now  represented  imperfectly  and  in  greatly  reduced  num- 
bers, but  the  areas  at  present  inhabited  by  their  descendants, 
except  in  the  case  of  the  Boreal  forms,  are  insignificant  in  com- 
parison with  their  former  extent.  It  should  be  remembered  that 
the  refrigeration  of  the  glacial  epoch  has  only  in  part  disappeared. 
In  earlier  Pliocene  times,  characteristic  representatives  of  sub- 
tropical faunas  and  floras  existed  northwards  over  much  of  the 
United  States  and  Canada,  and  in  still  earlier  times  reached  the 
Arctic  circle.  During  the  advance  of  cold  in  the  glacial  epoch 
these  forms  were  either  exterminated  or  driven  southward  into  the 
narrow  tropical  parts  of  Mexico  and  Central  America.  The  retreat 
of  cold  at  the  termination  of  this  period  was  not  complete,  and 
our  continent  has  never  regained  its  former  warmth.  Hence  the 
expelled  species  were  not  permitted  to  advance  more  than  a  short 
distance  into  the  region  formerly  occupied  by  them,  and  the 
tropical  species  have  been  held  back,  and  at  the  present  day  are 
not  found  except  along  the  extreme  southern  confines  of  our 
territory  [the  United  States].  For  example,  peccaries  in  early 
Plistocene  times  ranged  northward  over  a  large  part  of  western 
North  America,  while  at  present  they  are  restricted  to  parts  of 
Texas  and  Louisiana  below  the  Red  River  of  the  south  ;  and 
capivaras,  tapirs  and  other  tropical  forms  whose  fossil  remains 
have  been  found  in  many  parts  of  the  United  States  have  not  been 
able  to  return.  The  same  is  true  of  plants,  for  the  palms,  tree- 
ferns,  and  numerous  other  tropical  types  that  formerly  ranged  over 
much  of  our  country  are  now  either  altogether  extinct  or  exist 
only  in  the  tropics. 

"The  llama  and  many  plants  now  inhabiting  the  Andes  may 
be  looked  upon  as  representing  a  class  of  cases  in  which  Boreal 
forms  were  driven  so  far  south  that  they  actually  reached  the  great 
mountain-system  of  South  America  and  spread  southward  over  its 
elevated  plateaus  and  declivities  to  the  extreme  end  of  the  conti- 
nent in  Patagonia  and  Tierra  del  Fuego." 

1  Appendix,  No.  19,  p.  44. 


346 


THE    HOLARCTIC   REGION. 


[CHAP. 


Coming  to  the  consideration  of  sub-regions,  we  have  first  of 
all  the  Arctic  sub-region,  which  corresponds  to  the 
Boreal  sub-region  of  Dr  Heilprin,  and  the  Arctic 
zone  of  the  Boreal  region  of  Dr  Merriam,  and  is  of 
circumpolar  extent.  According  to  the  former  writer,  in  the  Old 
World  it  may  be  defined  as  the  tract  lying  to  the  north  of  a  line 
starting  from  about  the  66th  parallel  of  latitude  on  the  Norwegian 
coast,  and  passing  south-eastwards  to  the  coast  of  eastern  Asia 
in  about  the  5oth  parallel,  thus  including  the  greater  part  of 
Kamschatka,  and  Amurland.  In  America,  according  to  Dr 
Merriam's  map,  after  running  just  inside  the  shores  of  Newfound- 


FIG.  70.     MUSK-OX  (Ovibos  moschatus). 

land  and  Labrador,  the  boundary  line  bends  southwards  after 
passing  Cape  Chudleigh  to  coincide  with  the  southern  shore  of 
Hudson  Bay,  and  then  takes  a  north-westerly  direction  so  as 
to  include  within  the  sub-region  only  a  narrow  strip  on  the  north- 
eastern coast  of  Alaska,  and  a  somewhat  broader  one  on  the 
north-western  shore  of  the  same.  In  the  Old  World  the  boundary 
line  coincides  approximately  with  the  northern  limit  of  the  cultiva- 
tion of  cereals,  and  also  with  that  of  the  southern  migrations  of 
the  reindeer;  but  in  America  certain  reindeer  (which  are  regarded 


IX.]  ARCTIC   SUB-REGION.  347 

by  the  American  zoologists  as  specifically  distinct  from  thecircum- 
polar  "barren-ground"  variety)  extend  considerably  further  to  the 
south.  For  the  most  part  of  its  extent,  the  mammals  inhabiting 
this  sub-region  are  few  in  number,  a  large  proportion  of  them 
having  a  circumpolar  range.  Among  them  may  be  included  the 
Arctic  fox,  polar  bear,  wolverene,  the  ermine  or  stoat,  the  eastern 
and  western  species  of  lemming  (Myodes),  the  banded  lemming 
(Cuniculus  torquatus),  the  Arctic  vole  (Microtus  nitilus},  Parry's 
suslik  (Spermophilus  empetra],  the  musk-ox,  and  the  reindeer ; 
several  of  these  being  restricted  to  the  sub-region.  The  sea-otter 
(Latax)  frequents  the  shores  of  Alaska  and  Kamschatka,  but  also 
ranges  as  far  south  as  the  Kurile  Islands  and  California,  so  that  it 
is  not  confined  to  the  sub-region.  During  the  Plistocene  epoch, 
as  we  have  seen,  such  animals  as  the  mammoth,  horse,  bison,  and 
tiger  were  inhabitants  of  this  tract ;  the  latter  animal  being  still 
found  in  eastern  Siberia.  Towards  Amurland  and  the  Kams- 
chatkan  peninsula,  the  fauna  becomes  somewhat  less  scanty ;  the 
large  Kamschatkan  sheep  (Ovis  nivicola)  being  here  met  with,  as 
well  as  a  true  deer,  and  the  brown  bear. 

Of  other  groups  of  animals  inhabiting  the  more  typical  portions 
of  this  region,  there  may  be  noticed  among  the  birds  the  ptarmigan 
(Lagopus],  the  snowy  owl  (Nyctea  scandiaca),  the  Greenland  falcon 
(Falco  candicans),  the  eider-duck  (Somatcria  mollissima),  as  well  as 
various  species  of  divers  (Colymbus]  and  guillemots  (Uria  and 
Lomvia},  together  with  the  little  auk  (Mergulus  alle\  Dr  Heilprin 
writes  that  "  Captain  Markham  observed  the  footpiints  of  the 
polar  hare  in  the  snow-bound  ice  in  latitude  83°  10',  and  the 
antlers  of  a  reindeer  were  picked  up  by  the  officers  under  Sir 
George  Nares,  in  latitude  82°  45'  (Grinnell  Land).  A  skeleton  of 
the  latter  animal,  recently  picked  by  wolves,  was  also  obtained  in 
latitude  80°  27'.  Traces  of  the  rock-ptarmigan  (Lagopus  rupestris] 
have  been  met  with  as  far  north  as  latitude  83°  6',  and  the  snow- 
bunting  (Pkctrophanes  nivalis)  in  latitude  82°  33'.  The  reptile- 
fauna  is  very  limited,  no  serpent,  apparently,  passing  beyond  the 
sixty-seventh  parallel  of  latitude,  and  no  lizard  above  the  seventieth. 
The  fishes,  which  include  the  common  perch  and  pike,  are  mainly 
salmonoids.  Insects  are  fairly  numerous,  and  even  in  the  far 
north  the  number  of  species  is  considerable." 


348  THE    HOLARCTIC   REGION.  [CHAP. 

During  the  Plistocene  the  region  within  the  Arctic  circle  en- 
joyed a  decidedly  less  rigorous  climate  than  it  at  present  pos- 
sesses. In  Baron  von  Toll's  recent  expedition  to  the  New  Siberian 
Islands1,  where,  as  previously  stated,  remains  of  the  tiger  have  been 
obtained,  it  was  discovered  "  that  under  the  perpetual  ice,  in  a 
freshwater  deposit,  which  contained  pieces  of  willow  and  bones  of 
post-tertiary  mammals  (the  mammoth-layer)  were  complete  trees 
of  Alnus  fruticosa,  fifteen  feet  long,  with  leaves  and  fruit.  It  was 
thus  evident  that  during  the  mammoth-period  tree-vegetation 
reached  the  seventy-fourth  degree  of  latitude,  and  that  its  northern 
limit  was  at  least  three  degrees  further  north  than  it  is  now." 

The  next  sub-region  is  the  European,  which  may  be  taken  to 
include  all  that  part  of  Europe  lying  between  the 
Arctic  sub-region  in  the  north,  and  the  line  of  the 
Pyrenees  and  Alps,  continuing  eastwards  along  the 
northern  shore  of  the  Black  Sea  to  the  Caucasus  and  the  Caspian 
Steppes.  This  area  includes  the  typical  fauna  of  the  eastern 
Holarctic  region,  among  its  more  or  less  characteristic  mammals 
being  (in  the  north)  the  elk — also  ranging  into  America — ,  the  red 
deer  (unknown  in  America,  but  represented  by  a  variety  in  North 
Africa),  the  roe,  the  bison,  the  chamois,  the  Alpine  ibex,  the 
typical  variety  of  the  brown  bear,  the  badger,  the  wolverene  (in  the 
north),  the  Alpine  marmot  (Arctomys  marmotta),  the  dormouse, 
hamster,  mole,  and  hedgehog  \  several  of  these  being,  however, 
common  to  the  Arctic  and  Central  Asian  sub-regions.  The  des- 
mans (Myogale)  are  restricted  to  this  sub-region  ;  and  the  same 
was  probably  the  case  with  the  aurochs  (Bos  taurus,  var.  primi- 
genius],  the  ancestral  stock  of  our  domestic  cattle.  Finally,  the 
Caucasus  is  the  home  of  two  or  three  peculiar  species  of  goats 
(Capra  cylindricornis  and  C.  caucasica]  known  as  ture. 

It  will  be  unnecessary,  even  if  this  could  be  accomplished,  to 
give  a  complete  list  of  the  mammalian  fauna  of  this  sub-region, 
but  it  is  essential  to  refer  to  the  comparative  poverty  of  the  fauna 
of  the  British  Islands  as  compared  with  that  of  the  Continent. 
The  following  list  includes  all  the  mammals  (exclusive  of  bats) 
known  to  have  inhabited  the  British  Islands  within  the  historic 

1  See  Knowledge,  1895,  p.  106. 


IX.]  EUROPEAN   SUB-REGION.  349 

period ;  those  which  are  now  extinct  having  an  asterisk  prefixed 
to  them,  while  such  as  occur  in  Ireland  have  the  letter  I  added. 
Those  that  have  been  introduced  by  man  have  a  t  before  them. 
The  list  stands  as  follows,  viz. : — 

Hedgehog.     Erinaceus  europaeus.     I. 

Mole.     Talpa  europsea. 

Common  Shrew.     Sorex  araneus. 

Lesser          ,,  ,,      minutus.     I. 

Water-Shrew.     Crossopus  fodiens. 

Wild  Cat.     Felis  catus. 
*Wolf.     Canis  lupus.     I. 

Fox.  „     vulpes.     I. 

Pine-Marten.     Mustela  martes.     I. 

Polecat.  ,,        putorius. 

Stoat.  „        erminea.     I. 

Assogue.  ,,        hibernica.     I. 

Weasel.  ,,        vulgaris. 

Badger.     Meles  taxus.     I. 

Otter.     Lutra  vulgaris.     I. 
^Brown  bear.     Ursus  arctus.     I. 

Squirrel.     Sciurus  vulgaris.     I  (?  introduced), 
*  Beaver.     Castor  fiber. 

Dormouse.     Muscardinus  avellanarius. 

Harvest-Mouse.  Mus  minutus. 

Wood-Mouse.  ,,     sylvaticus.     I. 

Yellow-necked  Mouse.       ,,     flavicollis. 

Common  Mouse.  ,,     musculus.     I. 

f  Black  Rat.  „     rattus.     I. 

t  Brown  Rat.  ,,     decumanus.     I.. 

Common  Field:Vole.     Microtus  agrestis. 

Bank-Vole.  „         glareolus. 

Water-Vole  „         amphibius. 

Common  Hare.     Lepus  europaeus. 

Mountain  Hare.         ,,      timidus.     I. 
t  Rabbit.  ,,      cuniculus.     I. 

*?  Wild  Cattle.     Bos  taurus. 

Red  Deer.         Cervus  elaphus.     I. 


350  THE   HOLARCTIC    REGION.  [CHAP. 

t  Fallow  Deer.     Cervus  dama.     I. 

Roe  Deer.     Capreolus  caprea. 
*Wild  Boar.     Sus  scrofa.     I. 

The  total  number  in  this  list  is  only  28,  out  of  which  at  least 
four  are  introduced.  With  the  exception  of  the  recently-described 
assogue1,  which  is  intermediate  between  the  stoat  and  the  weasel, 
and  is  peculiar  to  Ireland,  the  whole  of  these  mammals  are  com- 
mon to  the  Continent.  As  shown  in  an  earlier  portion  of  the 
present  chapter,  during  the  Plistocene  epoch  Britain  possessed  a 
fauna  apparently  identical  with  that  of  the  Continent ;  and  there 
must  accordingly  be  some  good  reason  for  its  present  poverty  in 
mammalian  life  as  compared  to  the  latter  area.  The  difference  is 
accounted  for  by  Dr  Wallace,  through  the  occurrence  of  one  or 
more  periods  of  subsidence,  which  took  place  during  the  close  of, 
or  subsequent  to,  the  Glacial  epoch  ;  after  which  England  again 
became  united  to  the  Continent,  when  its  present  fauna  entered, 
the  period  of  connection  being,  however,  of  comparatively  short 
duration,  and  thus  permitting  of  the  passage  of  only  a  moiety  of 
the  continental  forms,  or  those  which  happened  at  the  time  in 
question  to  be  inhabiting  the  districts  nearest  to  the  connecting 
line.  Only  a  certain  number  of  the  mammals  which  thus  crossed 
into  Britain  have  ever  succeeded  in  reaching  Scotland ;  and  it  is 
from  this  country,  if  we  accept  the  views  of  Dr  Scharff,  referred  to 
above,  that  Ireland  appears  to  have  received  its  still  more  im- 
poverished mammalian  fauna. 

It  will  be  seen  that  the  foregoing  hypothesis  attributes  the 
clean  sweep  supposed  to  have  been  made  of  the  original  British 
fauna  to  the  effects  of  submergence,  and  not  to  the  ice-sheet.  On 
the  other  hand,  Mr  G.  W.  Bulman 2,  who  doubts  whether  the  sub- 
mergence has  been  sufficient  for  this,  attributes  such  extermination 
as  he  believes  to  have  taken  place  solely  to  the  effects  of  an  ice- 
sheet.  And  he  further  believes  that  a  number  of  the  original 
British  mammals  survived  in  the  southern  and  south-western 
counties  of  England,  whence  they  re-populated  Britain  on  the 
disappearance  of  the  ice-sheet,  without  there  having  been  any 

1  See  Thomas,  Natural  Science,  Vol.  vi.  p.  377  (1895). 
-  Appendix,  No.  12. 


IX.]  CENTRAL   ASIAN    SUB-REGION.  351 

subsequent  connection  with  the  Continent.  The  difficulty  con- 
nected with  this  explanation  is  that  it  apparently  necessitates  a 
pre-glacial  or  early  glacial  age  for  the  mammaliferous  deposits  of 
the  Thames  valley,  which  are  almost  certainly  inter-glacial  or 
post-glacial.  The  whole  subject  of  glaciation  is,  however,  so 
complicated  and  involved,  that  it  is  almost  impossible  to  form 
workable  theories  as  to  the  exact  mode  of  the  repopulation  of 
Britain  after  the  changes  which  took  place  during  the  glacial 
epoch. 

In  contrast  to  the  British  Isles,  which  are  eminently  of  the 
continental  type,  may  be  cited  Iceland,  lying  near  the  border-line 
between  the  Arctic  and  European  sub-regions,  which  is  as 
markedly  oceanic  in  its  character.  Beyond  an  occasional  ice- 
borne  polar  bear,  Iceland  possesses  only  the  Arctic  fox,  and  a 
mouse,  which  has  been  stated  to  be  a  peculiar  species ;  the  fox 
having  doubtless  been  originally  introduced  from  the  north  on 
floating  ice. 

According  to  the  scheme  of  Dr  Heilprin,  the  next  sub-region 
on  the  list  is  that  of  Central  Asia,  which  includes 
the  countries  bounded  on  the  west  by  the  European,  sSb-regi™*1*11 
and  on  the  north  by  the  Arctic  sub-region,  and 
extends  eastwards  as  far  as  Mantchuria  and  China  proper,  being 
bordered  on  the  south  by  the  Kuenlun  and  Nanshan  mountains1. 
A  large  portion  of  the  western  districts  of  this  tract  are  open 
steppes  or  deserts ;  and  in  such  tracts  several  peculiar  types  of 
rodents,  such  as  the  Kirghiz  jerboa  (Alactaga)  and  the  Yarkand 
jerboa  (Euchoretes\  are  met  with,  while  the  saiga  antelope  (Saiga), 
and  the  Mongolian  gazelle  ( Gazella  guttu rosd)  are  likewise  charac- 
teristic types.  Susliks  (Spermophilus),  marmots  (Arctomys),  and 
picas  are  very  abundant ;  and  the  place  of  the  European  wild  cat 
is  occupied  by  Pallas's  cat  (Felis  manul),  the  tiger  being  also 
sparingly  found  in  the  western  districts,  where  the  ounce  is  like- 
wise met  with.  In  part  of  this  sub-region  the  red  deer  is  replaced 
by  a  variety  or  species  known  as  Cervus  xanthopygus,  while 
Yarkand  is  the  home  of  a  variety  of  the  Kashmir  stag  (C.  cash- 
mirianus],  and  the  Thian-Shan  possesses  the  very  fine  and  wapiti- 

1  Dr  Heilprin  included  the  Tibetan  plateau  in  this  sub-region. 


352  THE    HOLARCTIC   REGION.  [CHAP. 

like  form  described  under  the  name  of  C.  eustephanus ;  all  these 
deer  being  mostly  inhabitants  of  forest-districts.  The  Tatarian 
roe  (Capreolus  pygargus),  inhabiting  suitable  localities  in  the  moun- 
tains forming  the  watershed  between  the  Russian  and  Chinese 
empires  and  Turkestan,  is  also  generally  regarded  as  specifically 
distinct  from  its  western  ally.  The  sub- region  is  also  the  chief 
home  of  the  magnificent  sheep  known  as  argali,  among  which 
the  splendid  Pamir  sheep  (Ovis  poll)  ranges  from  the  Pamirs  to 
the  Altai,  while  the  true  argali  (O.  ammon) — if  the  Tibetan  O.  hodg- 
soni  be  really  distinct — is  also  restricted  to  this  sub-region,  where 
it  is  now  confined  to  northern  Mongolia,  although  it  formerly 
inhabited  the  Altai.  The  ibex  of  the  Altai  is,  however,  identical 
with  the  Himalayan  and  Tibetan  Capra  sibirica. 

Although  included  by  Dr  Heilprin  in  the  preceding,  the  area 
typified  by  the  Tibetan  plateau  is  regarded  by  Dr 
Blanford '  as  constituting  a  sub-region  by  itself. 
Typically  this  region  is  bounded  on  the  north  by 
the  ranges  of  the  Kuenlun,  Altyn  Tag,  and  Nanshan,  and  extends 
eastwards  to  China  proper,  while  to  the  west  it  must  be  taken  to 
include  Ladak  and  the  upper  Indus  valley  as  far  as  Gilgit2.  To 
the  south  it  extends  to  the  main  chain  of  the  Himalaya.  The 
following  list  of  mammals  is  given  by  Dr  Blanford  as  distinctive  of 
this  sub-region ;  the  names  of  such  species  and  genera  as  are 
entirely  or  mainly  confined  to  the  area  being  printed  in  italics. 

Insect!  vora. 

Crocidura  aranea. 
Nectogale  elegans. 

Carnivora. 

Paradoxurus  laniger. 
Canis  lupus,  var.  laniger. 

,,      vulpes,  var.  flavescens. 

,,     ferrilatus. 

,,      deccanensis,  var. 
Mustek  foina,  var. 

,,         larva ta, 

1  Proc.  Zool.  Soc.  1893,  p.  449. 

2  See  Blanford,  Fauna  of  British  India,  Mammalia,  p.  v. 


IX.]  TIBETAN   SUB-REGION.  353 

Carnivora  (cont.\ 

Mustela  canignla. 

„         alpina,  var.  temon. 

„         erminea. 
Meles  leucura. 

„      albogularis. 
sEluropus  melanoleuctts. 
Ursus  pruinosus. 

Rodentia. 

Eupetaurus  rinereus. 
Arctomys  himalayanus. 

„         robustus. 
Mus  sublimis. 
Microtus  blythi. 
,,         strauchi. 
„        przevalskii. 
Siphneus  fontanieri. 
Lagomys  curzonice. 

,,        rutilus. 

,,        erythrotis. 

„        melanostomus . 

,,        ladacensis. 
Lepus  oiostolus. 
,,       hypsibius. 

Ungulata. 

Equus  hemionus,  var.  kiang. 
Bos  grunniens. 
Ovis  hodgsoni. 

„     vignei,  var. 

,,     nahura. 
Capra  sibirica. 
Pantholops  hodgsoni, 
Budorcas  taxicolor. 
Gazella  picticaudata. 
Cervus  affinis. 

,,        thoroldi. 
Moschus  moschiferus. 
L.  23 


354  THE   HOLARCTIC   REGION.  [CHAP. 

This  list  includes  all  the  species  inhabiting  the  plateau  at 
elevations  exceeding  12,000  feet.  Dr  Blanford  writes  that  "many 
of  the  forms  named  only  inhabit  small  portions  of  the  area,  and 
whilst  Bos  grunniens,  Pantholops  hodgsoni,  and  Gazella  picti- 
caudata,  with  several  rodents,  appear  to  be  peculiar  to  the  high 
plateaus  above  14,000  feet,  the  two  species  of  Cervus  are  pro- 
bably found  in  brushwood  at  a  rather  lower  elevation  in  the  more 
broken  region  of  Eastern  Tibet,  where  the  rainfall  is  heavier  and 
the  vegetation  more  abundant. 

"As  was  printed  in  the  paper  in  the  Geological  Magazine1, 
there  is,  so  far  as  I  am  aware,  no  equally  peculiar  mammalian 
fauna  to  be  found  in  any  continental  area  of  equal  extent,  and  for 
a  parallel  it  is  necessary  to  turn  to  some  island  like  Celebes,  that 
has  long  been  isolated  from  all  surrounding  lands." 

This,  however,  is  not  all,  for  there  occur  at  Hundes,  on  the 
Tibetan  plateau,  mammaliferous  strata  yielding,  among  other 
remains,  bones  of  a  rhinoceros,  and  of  an  antelope  which  is 
apparently  generically  identical  with  the  chiru  (Pantholops),  now 
inhabiting  the  same  area.  The  isolation  and  development  of  this 
most  peculiar  fauna  is  intimately  connected  with  the  date  of 
elevation  of  the  Himalaya.  After  pointing  out  that  both  the  fossil 
chiru  and  the  fossil  rhinoceros  appear  to  have  inhabited  the  area 
when  it  had  attained  something  approaching  its  present  enormous 
elevation,  Dr  Blanford2  writes  as  follows  :  "  Bearing  in  mind  that 
the  isolation  of  the  Tibetan  plateau  is  far  less  perfect  as  regards 
mammals  than  that  of  any  island,  and  that  some  of  the  forms — 
the  Carnivora  especially — found  in  Tibet  are  evidently  very  recent 
immigrants,  it  is  a  reasonable  conclusion  that  the  peculiar  fauna 
of  the  Tibetan  plateau  has  been  distinct  from  that  of  neighbouring 
countries  since  middle  Tertiary  times. 

"  But  what  has  caused  the  isolation  of  the  Tibetan  fauna  ? 
Why  in  this  one  continental  tract  is  there  a  generic  and  specific 
differentiation  of  the  mammalia,  of  which  no  other  example 
exists  ?  There  is  only  one  character  in  which  Tibet  is  different 
from  other  continental  areas,  its  great  height.  This  alone  renders 
the  climate  of  Tibet  so  different  from  that  of  other  parts  of 

1  Decade  3,  Vol.  ix.  p.  161  (1892). 

2  Geol.  Mag.  op.  cit.  p.  165. 


IX.]  MANTCHURIAN   SUB-REGION.  355 

Central  Asia,  which  are  equally  cold  and  barren.  It  seems  a 
reasonable  inference  that  the  elevation  of  the  Tibetan  plateau 
dates  back  to  middle  Tertiary  times. 

"  It  is  of  course  probable  that  the  elevation  was  gradual ;  and 
although  the  area  may  have  been  sufficiently  high  at  the  close  of 
the  Miocene  period  to  produce  a  difference  in  climatal  conditions, 
the  greater  part  of  the  upward  movement  may  have  been  post- 
Miocene,  and  a  great  part  post-Pliocene." 

Bordering  as  it  does  upon  the  tropics,  where  it  abuts  against  the 
Oriental  region,  the  Mantchurian  sub-region  is  not 
easy  to  define,  since  the  intermingling  of  Holarctic 
and  Oriental  types  is  very  strongly  marked  on  its 
southern  confines.  Starting  somewhere  about  the  Amur  river,  it 
may,  however,  be  taken  to  include  the  Japanese  islands,  Mant- 
churia,  Corea,  and  northern  China ;  its  southern  limit  being  placed 
approximately  in  the  latitude  of  Fuchau.  Westwards  it  may  be 
taken  to  include  Moupin,  in  Eastern  Tibet,  although  this  district 
is  referred  by  Dr  Wallace  to  the  Oriental  region. 

From  all  the  other  sub-regions,  with  the  exception  of  the 
Mediterranean,  the  Mantchurian  is  distinguished  by  the  presence 
of  monkeys  belonging  to  the  genera  Macacus  and  Semnopithecus, 
some  of  these  occurring  in  Japan  and  others  in  Eastern  Tibet. 
Of  the  latter,  one  (Semnopithecus  roxellance)  is  peculiar,  and  the 
other  is  identified  by  Mr  H.  O.  Forbes  with  the  widely-spread 
Oriental  Macacus  arctoides.  Among  the  Carnivora,  the  Oriental 
genus  Helictis  enters  this  sub-region,  one  species  occurring  in  the 
neighbourhood  of  Shanghai ;  while  Japan  is  the  home  of  a  peculiar 
long-haired  dog  ( Cants  procyonides),  which  is  frequently  separated 
generically  under  the  name  of  Nyctereutes,  although  it  unquestion- 
ably pertains  to  the  typical  genus.  Perhaps,  however,  the  most 
characteristic  mammals  are  the  deer.  Foremost  among  these  are 
a  group  of  small  deer  belonging  to  the  genus  Cenms,  and  distin- 
guished from  the  red  deer  group  by  the  invariable  absence  of  a 
bez-tine  to  the  antlers,  each  of  which  has  but  four  points.  These 
deer  are  further  characterised  by  the  coat  of  the  adult  being 
spotted  in  summer  with  white,  but  uniformly  brown  in  winter,  and 
also  by  the  black  lateral  margins  to  the  white  blaze  on  the  hind- 
quarters. The  species  include  the  Japanese  deer  (C.  sica), 

23—2 


356  THE   HOLARCTIC   REGION.  [CHAP. 

common  to  Japan  and  North  China,  the  larger  Mantchurian  deer 
(C.  mantchuricus\  and  Dybowski's  deer  (C.  dybowskii)  from  the 
upper  Ussuri  district  of  Mantchuria,  in  the  neighbourhood  of 
Vladivostock.  Elsewhere  the  group  is  represented  in  Formosa, 
and  also  in  the  Caspian  provinces  of  Persia.  In  addition  to 
these,  there  are  the  hornless  Chinese  water-deer  (Hydropoles),  and 
the  two  species  of  tufted  deer  (Elaphodus)\  the  latter  being  closely 
allied  to  the  Oriental  muntjacs.  What  is  known  of  the  palseonto- 
logical  history  of  the  southern  portion  of  this  area  indicates  that 
during  the  Pliocene  epoch  its  mammalian  fauna  was  closely  allied 
to  that  of  the  Siwalik  Hills,  thus  showing  that  at  this  time  there 
was  no  distinction  between  the  Oriental  and  Holarctic  regions, 
which  even  now  grade  imperceptibly  into  one  another  in  this 
district. 

The  remains  of  fossil  elephants  from  Japan1  are  referable  to 
Elephas  clifti,  insignis,  and  namadicus,  of  which  the  two  first  are 
common  to  the  Siwaliks,  while  the  third  occurs  typically  in  the 
Plistocene  Narbada  beds  of  India.  From  the  known  distribution 
of  these  elephants,  it  is  probable  that  Japan  was  connected  with 
the  mainland  during  the  Pliocene  epoch  by  way  of  the  Corean 
peninsula,  although  Dr  Wallace  is  of  opinion  that  its  latest  con- 
nection was  to  the  north.  Of  existing  animals  common  to  Japan 
and  the  mainland,  allusion  has  already  been  made  to  Cervus  sica  : 
and  another  common  type  is  the  giant  salamander  Megalobatra 
chus.  The  latter  genus  is  represented  in  a  fossil  state  in  the 
Miocene  of  Baden,  and  as  it  is  closely  allied  to  the  North  American 
CryptobranchuS)  there  is  clear  evidence  of  the  eastern  migration  of 
this  ancient  type,  of  which  the  two  survivors  are  respectively  con- 
fined to  China  and  Japan  on  the  one  hand,  and  North  America 
on  the  other.  Further  evidences  of  affinity  between  the  fauna  of 
Japan  and  North  America  are  afforded  by  the  circumstance  that 
one  species  of  the  mole-like  genus  Urotrichus  is  confined  to  the 
former  islands,  while  the  other  is  an  inhabitant  of  the  north- 
western districts  of  the  latter  continent.  The  sea-otter  (Latax]  is 
likewise  common  to  the  coasts  of  Japan,  the  Kurile  Islands,  and 
Kamschatka,  and  the  Pacific  shores  of  North  America.  More 
remarkable,  however,  is  the  fact  that  a  North  American  scincoid 
1  See  Naumann,  Palceontographica,  Vol.  xxvm.  Art.  r  (1881). 


IX.]  MEDITERRANEAN    SUB-REGION.  357 

lizard  (Eumeces  quinquelineatus]  is  represented  in  Japan  by  a  form 
(E,  marginatus]  so  closely  allied  that  the  two  were  long  considered 
inseparable,  although  they  are  now  regarded  as  distinct1.  All 
these  facts  are  indicative  that  Japan  was  formerly  joined  to  both 
Corea  and  Kamschatka,  whence  land  was  continued  across  Bering 
Strait  to  unite  the  Old  World  with  Alaska. 

Although,  as  already  stated,  the  Mediterranean  or  Tyrrhenian 
sub-region   has    strong   claims    to    be   regarded   as        Mediterra 
representing  a  region  by  itself,  it  may  be  more  con-     nean  Sub- 
veniently   considered   here   than    later    on    in   the 
chapter.     In  addition  to  such  parts  of  Africa  and  Arabia  as  lie 
to  the  north  of  the  Ethiopian  region,   this  sub-region  includes 
Spain,  those  parts  of  Europe  situated  south  of  the  Alps,  together 
with  Turkey,  Asia  Minor,  Persia,  Baluchistan,  and  Afghanistan. 
Whether  Kashmir  should  be  regarded  as  an  aberrant  outlier  of 
this  region,  I  am  not  yet  satisfied.     Although  gerbils  (Gerbillus*) 
are   also   found   in    the    Oriental   and    Ethiopian    regions,    their 
distribution  in  the  Holarctic  is  very  nearly  coincident  with  the 
limits  of  the  present  sub-region. 

Whereas  to  the  north  of  the  Mediterranean  Sea  a  large  pro- 
portion of  the  mammals  are  more  or  less  typically  Holarctic,  in 
North  Africa  and  Syria  those  with  an  Ethiopian  facies  are  met 
with,  and  an  Oriental  element  makes  its  appearance  in  the  eastern 
districts  of  the  sub-region.  Even  in  Africa,  however,  some  of  the 
forms  have  an  Oriental  facies,  the  Barbary  ape  (Macacus  inuus) 
belonging  to  a  genus  whose  home  is  now  in  the  Oriental  region, 
and  which  is  totally  unknown  in  the  Ethiopian.  As  a  wanderer 
from  the  Ethiopian  region,  mention  may  first  be  made  of  a  species 
of  jumping -shrew  (Macroscelides)  met  with  in  Barbary,  while 
among  the  octodont  family  of  the  rodents,  the  gundi,  forming 
the  sole  representative  of  the  genus  Ctenodactylus,  has  its  nearest 
allies  in  Ethiopia,  although  it  is  confined  to  North  Africa.  The 
Barbary  ape,  although  occurring  on  the  rock  of  Gibraltar,  where  it 
may  have  been  introduced,  is  otherwise  confined  to  North  Africa. 

1  See  Boulenger,  Cat.  Lizards,  Brit.  Mtis.  Vol.  III.  p.  369. 

2  Many  writers  separate  certain  species  as  Meriones,  but  as  the  two  groups 
are  connected  by  G.  indicus  (see  Lataste,  Proc.  ZooL  Soc.  1884,  p.  88),  such 
distinction  seems  superfluous. 


358  THE   HOLARCTIC   REGION.  [CHAP. 

In  the  Carnivora,  the  striped  hyaena,  which  is  also  an  inhabitant 
of  India,  is  widely  spread  in  this  sub-region,  ranging  through 
western  Asia  to  northern  Africa.  The  common  genet  (Genetta 
vulgaris\  which  belongs  to  a  genus  otherwise  exclusively  Ethio- 
pian, is  mainly  confined  to  this  region,  inhabiting  southern  France, 
Spain,  Turkey,  North  Africa,  and  Palestine.  A  nearly  similar 
distribution  characterises  the  common  mungoose,  or  ichneumon 
(Herpestes  ichneumon],  which  frequents  southern  Spain,  Asia 
Minor,  North  Africa,  and  Palestine.  The  large  weasel  (Mustela 
africanus)  common  to  Egypt,  Malta,  and  perhaps  the  south  of 
Italy  has  been  already  referred  to  in  an  earlier  part  of  this  chapter. 
In  addition  to  Ctenodactylus,  the  rodents  possess  another  and  more 
widely-spread  generic  type  confined  to  this  sub-region  in  the  form 
of  the  great  mole-rat  (Spalax  typhlus),  whose  range  includes  south- 
eastern Europe,  Persia,  Mesopotamia,  Syria,  and  Egypt.  In  the 
same  order  the  common  porcupine  (ffystrix  cristata],  although 
ranging  into  West  Africa,  is  found  but  little,  if  at  all,  to  the  north 
of  the  present  sub-region,  where  it  is  common  to  northern  Africa 
and  southern  Europe. 

Among  the  ungulates  the  addax  antelope  (Addax  nasomacu- 
lata\  although  allied  to  Ethiopian  types,  is  solely  Mediterranean, 
its  home  being  North  Africa  and  Syria.  More  closely  allied  to 
the  Ethiopian  fauna  are  certain  hartebeests  of  the  genus  Bubalis, 
the  smaller  of  which  (B.  mauritanica]  is  common  to  North  Africa, 
Syria,  and  Arabia,  while  the  second  {B,  major]  inhabits  Tunis. 
The  same  is  the  case  with  the  Beatrix  antelope  (Oryx  beatrix] 
of  Western  Arabia  and  Bushire.  In  gazelles,  this  sub-region  is 
remarkably  rich,  doubtless  from  the  number  of  sandy  or  desert 
tracts  it  contains.  Algeria  is  the  habitat  of  the  three  species 
known  as  Gazella  loderi,  G.  kevella,  and  G.  rufina,  while  G. 
dorcas  ranges  through  Egypt,  Algeria,  Syria,  Palestine,  and  a  part 
of  Asia  Minor,  and  G.  subgutturosa  roams  from  Persia  through 
Afghanistan  and  Turkestan.  The  aberrant  sheep  known  as  the 
arui  (Ovis  tragelaphus]  is  now  restricted  to  North  Africa;  and  the 
mouflon  (O.  musimon),  although  its  fossil  remains  have  been  found 
on  the  Continent,  appears  to  be  now  restricted  to  Corsica.  An- 
other species  peculiar  to  the  sub-region  is  the  Armenian  sheep 
(O.  gmelini)  of  eastern  Persia  and  Asia  Minor,  represented  by  a 


IX.]  KASHMIR.  359 

closely-allied  form  in  Cyprus.  Of  the  goats,  the  Spanish  ibex 
(Capra  pyrenaica)  is  restricted  to  the  mountains  of  Spain;  while 
the  Sinaitic  ibex  (C.  sinaitica)  represents  the  group  in  Palestine 
and  upper  Egypt.  Among  the  Cervidcz,  the  two  species  of  fallow- 
deer  were  originally  confined  to  this  area,  the  common  Cervus 
dama  being  a  native  of  the  Mediterranean  countries,  while  the 
Persian  C.  mesopotamicus  is  found  in  the  mountains  of  Luristan,  in 
Mesopotamian  Persia.  In  North  Africa  the  ordinary  red  deer  is 
represented  by  a  variety  distinguished  by  the  invariable  absence  of 
a  bez-tine  to  the  antlers.  A  connection  with  the  Tibetan  sub- 
region  is  afforded  by  the  wild  asses  inhabiting  the  desert-plains 
between  the  Red  Sea  and  the  Indus,  since  both  these  and  the 
Tibetan  form  are  but  varieties  of  a  single  species  (Equus  hemi- 
onus).  Lastly,  Ethiopian  affinities  are  exhibited  by  the  occurrence 
of  a  species  of  hyrax  (Procavid]  in  Syria.  In  the  early  part  of  the 
present  century  the  hippopotamus  still  inhabited  lower  Egypt, 
while,  as  we  have  seen,  the  lion,  which  is  now  common  in  parts  of 
Persia,  was  found  within  the  historic  period  in  Thrace.  At  a  still 
earlier  date,  both  these  animals,  as  well  as  the  spotted  hyaena, 
extended  as  far  north  as  England. 

On  the  whole,  therefore,  the  fauna  of  this  sub-region  is  a  very 
mixed  one ;  and  this  fact,  together  with  the  difficulty  in  defining 
its  boundaries,  suggests  the  need  of  further  deliberation  before  the 
area  is  raised  to  the  rank  of  a  separate  region.  The  former  con- 
nections between  southern  Europe  and  Africa  having  been  alluded 
to  in  an  earlier  part  of  the  chapter,  require  no  further  notice  in 
this  place. 

Very  difficult  to  determine  is  the  position  which  should  be 
given  to  the  valley  of  Kashmir,  since  its  fauna 
exhibits  such  a  mingling  of  Oriental  and  Holarctic 
types  that  it  might  almost  be  as  well  assigned  to  one  region  as  the 
other.  Holarctic  affinities  are,  however,  exhibited  by  the  occur- 
rence of  a  species  of  the  red  deer  group,  Cervus  cashmirianus,  and 
likewise  by  one  of  the  rodent  genus  Sminthus,  of  which  the 
second  species  inhabits  northern  and  eastern  Europe,  and  the 
third  Kansu,  in  western  China.  A  variety  of  the  brown  bear  is 
also  indicative  of  Holarctic  affinities,  and  this  is  still  more 
markedly  the  case  with  the  spiral-horned  goat  known  as  the 


360  THE    HOLARCTIC    REGION.  [CHAP. 

markhor  (Capra  falconeri],  of  which  one  variety  inhabits  the  Pir 
Panjal  range,  on  the  south  side  of  the  valley,  while  the  others  are 
found  in  the  districts  to  the  north  and  west  of  Kashmir.  The 
musk-deer,  again,  is  another  essentially  Holarctic  type.  On  the 
other  hand,  the  occurrence  of  a  langur -(Semnopithecus)  and  a  ma- 
caque (Macacus)  points  to  a  connection  with  the  Oriental  fauna ; 
and  a  Kashmir  mungoose  (Herpestes  auropunctatus]  is  identical 
with  one  from  India.  There  are,  however,  none  of  the  exclusively 
Oriental  genera  in  Kashmir ;  and  this  fact,  coupled  with  the 
absence  of  all  deer  of  the  sambar-group,  leaves  little  doubt  that 
the  valley  really  belongs  to  the  Holarctic.  Whether  it  should 
be  regarded  as  pertaining  to  the  Mediterranean  sub-region,  or  as 
forming  a  distinct  sub-region  by  itself,  must  be  reserved  for  future 
consideration. 

Passing  to  the  western  division  of  the  Holarctic  region,  the 
tract  lying  to  the  southward  of  the  circumpolar  Arctic 
sub-region,  designated  by  Dr  Merriam  the  Boreal 
zone  of  his  Boreal  region,  may  be  conveniently 
termed  the  Canadian  sub-region.  Its  northern  boundary  is,  of 
course,  identical  with  the  southern  limits  of  the  Arctic  sub-region, 
and  the  area  includes  the  greater  part  of  the  Dominion  of  Canada, 
although  a  long  strip  runs  down  the  line  of  the  Rocky  Mountains, 
and  another  along  the  Pacific  coast,  into  the  United  States.  Indeed, 
Dr  Merriam  includes  in  this  sub-region  all  the  higher  plateaus  of 
Wyoming  and  Colorado,  so  that  the  sub-region  embraces  a 
number  of  small  disconnected  areas  on  its  south-western  ex- 
tremity, and  it  is  consequently  impossible  to  define  its  limits  by 
description.  It  may  be  stated,  however,  that  on  the  eastern  side 
of  the  continent  the  sub-region  extends  from  Hudson  Bay  to  the 
middle  of  Lake  Michigan,  while  on  the  western  coast  it  stretches 
from  near  the  extremity  of  Alaska  to  San  Francisco ;  a  big  loop 
extending  northwards  of  Montana  nearly  to  latitude  55°. 

The  mammalian  fauna  of  the  Canadian  sub-region  is  that  of 
the  western  division  of  the  Holarctic  region  generally,  and  includes 
those  forms  mentioned  on  page  344.  According  to  Dr  Merriam, 
the  following  genera  from  this  sub-region  do  not  range  further 
south  than  the  undermentioned  Transition  zone ;  namely,  Condy- 
lura,  Urotrichus,  Gulo,  Latax,  Arctomys,  Haplodon,  Phenacomys, 


IX.]  TRANSITION   ZONE.  361 

Myodes,  Cuniculus,  Zapus,  Erethizon,  Lagomys,  Cervus,  Alces, 
Rangifer,  and  Haploceros.  On  the  other  hand,  the  following, 
which  are  as  clearly  of  northern  origin,  penetrate  as  far  south 
as  the  Sonoran  region,  which  some  of  them  enter.  These  are 
Sorex,  Mustela  (only  the  members  of  the  sub-genus  Putorius), 
Ursus,  Fiber,  Microtus,  Castor,  Tamias,  Bos,  and  Ovis. 

Between   the  Canadian   sub-region  of  the  Holarctic  and  the 
Sonoran  region  is  interposed  a  tract  whose  fauna 
contains  a  mixture  of  Canadian  and  Sonoran  forms,        Transition 

Zone. 

and  it  is  consequently  termed  by  Dr  Merriam  the 
Transition  zone.  Under  this  somewhat  indefinite  title  the  area 
may  best  be  left.  It  is  described  by  the  author  just  cited  as 
follows1:  "The  humid  division  of  this  zone,  known  as  the 
Alleghanian  fauna,  covers  the  greater  part  of  New  England  (except 
Maine  and  the  mountains  of  Vermont  and  New  Hampshire),  and 
extends  westerly  over  the  greater  part  of  New  York,  southern 
Ontario,  and  Pennsylvania,  and  sends  an  arm  south  along  the 
Alleghanies,  all  the  way  across  the  Virginias,  Carolinas,  and 
eastern  Tennessee,  to  northern  Georgia  and  Alabama.  In  the 
Great  Lake  region  this  zone  continues  westerly  across  southern 
Michigan  and  Wisconsin,  and  then  curves  northward  over  the 
prairie-region  of  Minnesota,  covering  the  greater  parts  of  North 
Dakota,  Manitoba,  and  the  plains  of  the  Saskatchewan ;  thence 
bending  abruptly  south,  it  crosses  eastern  Montana  and  Wyoming, 
including  parts  of  western  South  Dakota,  and  Nebraska,  and  forms 
a  belt  along  the  eastern  base  of  the  Rocky  Mountains  in  Colorado 
and  northern  New  Mexico,  here  as  elsewhere  occupying  the 
interval  between  the  Upper  Sonoran  and  Canadian  zones. 

"  In  Wyoming  the  Transition  zone  passes  broadly  over  the 
well-known  low  divide  of  the  Rocky  Mountains,  which  affords  the 
route  of  the  Union  Pacific  railway,  and  is  directly  continuous  with 
the  same  zone  in  parts  of  Colorado,  Uta,  and  Idaho,  skirting  the 
Canadian  boundaries  of  the  Great  Basin  all  the  way  around  the 
plains  of  the  Columbia,  sending  an  arm  northward  over  the  dry 
interior  of  British  Columbia,  descending  along  the  eastern  base  of 
the  Cascade  Range  and  the  High  Sierra  to  the  southern  extremity 

1  Appendix,  No.  19,  p.  30.     In  this  extract  the  word  Canadian  has  been 
substituted  for  Boreal. 


362  THE   HOLARCTIC   REGION.  [CHAP.  IX. 

of  the  latter,  and  occupying  the  summits  of  the  Coast  Ranges  in 
California  and  of  many  of  the  desert  ranges  of  the  Great  Basin. 

"The  Transition  zone,  as  its  name  indicates,  is  a  zone  of 
overlapping  of  Canadian  and  Sonoran  types.  Many  Canadian 
genera  and  species  here  reach  the  southern  limits  of  their  distribu- 
tion, and  many  Sonoran  genera  and  species  their  northern  limits. 
But  a  single  mammalian  genus  (Synaptomys)  is  restricted  to  the 
Transition  zone,  and  future  research  may  show  that  it  inhabits  the 
Canadian  region  also." 


FIG.  71.     MUSQUASH  (Fiber  zibet  hi  cus). 

The  writer  adds,  however,  that  there  are  a  considerable  number 
of  species — mostly  rodents — restricted  to  this  zone.  The  follow- 
ing Canadian  genera,  namely,  Condylura,  Urotrichus,  Ursus, 
Arctomys,  Tamias,  Fiber1,  Zapus,  Erethizon,  Cervus,  and  Ovis, 
almost  or  completely  disappear  in  this  zone ;  while  the  following 
intruders  from  the  Sonoran,  namely  Scalops,  Bassariscus,  Spilo- 
gale,  Perognathus,  Thomomys,  Geoitiys,  Cynomys  and  Antilocapra 
do  not  range  further  north,  several  of  them,  indeed,  only  intruding 
into  the  zone  in  a  small  area  in  the  west. 

1  Penetrates  the  Sonoran  along  the  lines  of  streams  where  cool  currents  of 
air  are  carried  down. 


CHAPTER   X. 
THE  SONORAN  REGION. 

Limits — Characteristics  of  Mammalian  Fauna — Extinct  Groups  of  Mammals 
characteristic  of  Western  Arctogaea — Distinctness  of  the  Region — Dual 
Origin  of  Groups. 

As  stated  in  the  introductory  chapter,  wherever  one  zoological 
region  of  the  globe  has  no  definite  physical  barrier  by  which 
it  is  separated  from  the  next  well-marked  region,  there  must 
always  occur  an  intermediate  tract  where  the  characteristic  types 
of  the  faunas  of  the  two  regions  inosculate  and  intermingle.  That 
this  is  the  case  with  that  area  of  North  America  denominated  the 
Sonoran  region  has  been  indicated  at  the  close  of  the  preceding 
chapter,  and  the  existence  of  the  Transition  zone,  which  seems, 
on  the  whole,  to  pertain  to  the  Holarctic  region,  unfortunately 
prevents  the  Sonoran  from  being  defined  with  the  precision  which 
would  be  possible  had  this  area  a  high  mountain-barrier  on  its 
northern  frontier. 

In  a  map  of  the  small  dimensions  of  the  one  accompanying 
this  volume  it  is  impossible  to  show  with  any 

Limits. 

attempt  at  accuracy  the  complex  nature  of  the 
northern  boundary  of  this  region,  which  will,  however,  be  found 
accurately  laid  down  in  the  map  illustrating  Dr  Merriam's  memoir1. 
According  to  the  latter  writer,  "  the  Sonoran  region  as  a  whole 
stretches  across  the  continent  from  the  Atlantic  to  the  Pacific, 
covering  nearly  the  whole  country  south  of  latitude  43°,  and 
reaching  northward  on  the  Great  Plains  and  Great  Basin  to  about 
latitude  48°.  It  is  invaded  from  the  north  by  three  principal 
intrusions  of  Canadian2  forms  along  the  three  great  mountain- 

1  Appendix,  No.  19. 

2  Boreal  in  the  original. 


364  THE   SONORAN    REGION.  [CHAP. 

systems  already  mentioned  [Alleghanies,  Rocky  Mountains,  and 
Cascade  and  Sierra  Nevada  ranges] ;  while  to  the  southward  it 
occupies  the  great  interior  basin  of  Mexico,  and  extends  into  the 
tropics  along  the  highlands  of  the  interior.  It  covers  also  the 
peninsula  of  lower  California,  the  southern  part  of  which  seems 
entitled  to  rank  as  an  independent  subdivision." 

Later  it  is  stated  that  the  region  "may  be  divided  by  tempe- 
rature into  two  principal  transcontinental  zones,  Upper  Sonoran 
and  Lower  Sonoran ;  and  each  of  these  in  turn  may  be  subdivided 
into  arid  and  humid  divisions." 

The  proposal  to  form  a  separate  region  for  such  an  insignifi- 
cant area  as  the  southern  extremity  of  California  seems  unnecessary, 
although  its  fauna  may  differ  considerably  from  that  of  the  typical 
Sonoran. 

Omitting  mention  of  the  bats,  the  mammalian  genera  charac- 

Characteris       teristic  of  the  Sonoran  region  may  now  be  taken  into 

tics  of  Mam-       consideration.     Commencing  with  the  Insectivora, 

tnalian  Fauna.         ,  ,  .. 

the  Sonadcz  are  represented  by  the  peculiar  genus 
Notiosorex,  which  is  closely  allied  to  the  Oriental  Soriculus, 
but  has  only  28  in  place  of  30  teeth.  Of  this  genus  the  two 
species  do  not  range  north  of  this  region,  although  they  also 
enter  Central  America1.  The  short -tailed,  or  earless  shrews 
(Blarina),  with  either  32  or  30  teeth,  are  also  mainly  Sonoran, 
although  ranging  northwards  into  the  Holarctic,  and  southwards 
into  Guatemala.  In  the  Talpida  the  three  species  of  the  mole-like 
genus  Scalops,  characterised  by  having  36  teeth,  webbed  hind  feet, 
and  a  short  and  nearly  naked  tail,  are  mainly  Sonoran,  although 
passing  into  the  Transition  zone.  On  the  other  hand,  the  two 
species  of  Scapanus,  distinguished  by  the  possession  of  40  teeth, 
and  the  hairy  tail,  have  a  distribution  very  similar  to  Blarina, 
although  they  do  not  enter  Central  America. 

In  the  Carnivora  the  raccoon-family  (Procyonidce)  is  very 
strongly  represented,  although  none  of  the  genera  are  absolutely 
peculiar  to  the  region.  The  genus  Bassariscus — a  near  ally  of  the 
true  raccoons,  and  possessing  two  species — is  nevertheless  mainly 
Sonoran,  although  it  ranges  into  the  Transition  zone  of  the 

1  Teste  Dobson. 


X.]  ITS    FAUNA.  365 

Holarctic  and  also  into  Central  America.  The  true  raccoons,  on 
the  other  hand,  cannot  be  regarded  as  distinctive  of  the  region, 
since  they  range  from  South  America  into  the  Canadian  sub- 
region  of  the  Holarctic ;  and  the  coatis  (Nasua)  are  now  highly 
characteristic  of  the  Neogaeic  realm.  Indeed  Dr  Merriam  con- 
siders both  genera  as  intruders  from  the  latter  realm,  but  this  can 
scarcely  be  regarded  as  the  correct  view.  The  family  is  repre- 
sented in  the  two  halves  of  the  northern  hemisphere  (in  the  eastern 
by  sElurus],  in  both  of  which  it  dates  from  the  Pliocene,  and, 
as  it  is  unknown  in  South  America  till  the  Plistocene  or  late 
Pliocene,  it  is  evidently  one  of  northern  origin ;  the  American 
forms  having  probably  attained  their  maximum  development  in 
the  Sonoran  region.  Much  the  same  is  the  case  with  regard  to 
the  skunks  among  the  Mustelidcz ;  these  being  probably  an  original 
Sonoran  type  which  has  spread  northwards  into  the  Holarctic 
region  and  southwards  into  the  Neogaeic  realm.  Of  these,  the 
single  species  of  climbing  skunk  (Sptlogale)  is  mainly  Sonoran, 
although  it  also  enters  the  Transition  zone  of  the  Holarctic,  and 
likewise  Central  America.  Of  the  other  members  of  the  group, 
the  typical  skunks  (Mephitis)  range  from  Hudson  Bay  to  Guate- 
mala ;  while  the  allied  genus  Conepatus  is  found  from  Texas  to 
Patagonia.  In  the  same  family  the  American  badgers  (Taxided), 
although  ranging  well  into  the  Holarctic,  are  regarded  by  Dr 
Merriam  as  of  Sonoran  origin.  These  badgers,  it  may  be  ob- 
served, differ  from  the  true  badgers  of  the  Old  World  by  the  form 
and  characters  of  their  cheek-teeth,  the  last  upper  molar  being 
proportionately  much  smaller. 

Turning  to  the  rodents,  the  well-known  prairie -marmots 
( Cynomys],  which  occupy  a  position  intermediate  between  the  true 
marmots  and  susliks,  are  regarded  by  Dr  Merriam  as  of  Sonoran 
origin,  although  they  extend  into  the  Holarctic.  In  the  Muridce 
the  peculiar  cricetine  genus  Rhithrodontomys — which,  together  with 
the  allied  South  American  Rhithrodon,  differs  from  the  other 
members  of  the  sub-family  to  which  it  belongs  by  its  grooved 
upper  incisors — appears  to  be  restricted  to  the  region  under  con- 
sideration. The  white-footed  mice  (Sitomys),  although  distributed 
over  the  whole  of  the  New  World,  seem  to  attain  their  maximum 
specific  development  in  the  Sonoran,  to  which  the  two  sub-genera 


366  THE   SONORAN    REGION.  [CHAP. 

Onychomys  and  Oryzomys  are  restricted.  Yet  their  near  alliance 
to  the  Old  World  hamsters  indicates  that  the  group  must  have 
had  a  northern  origin,  although  the  genus  may  have  attained 
its  present  distinctive  features  within  the  Sonoran  area.  The 
genus  Sigmodon,  which  differs  from  the  last  in  the  pattern  of  the 
molar  teeth,  and  is  represented  solely  by  the  rice-rat,  does  not 
range  north  of  the  Sonoran  region,  although  extending  into  South 
America  as  far  as  Ecuador.  The  wood-rats  (Neotoma),  in  which 
the  molars  simulate  the  prismatic  appearance  of  those  of  the  voles, 
.are  also  largely  Sonoran,  although  they  extend  into  the  Canadian 


FIG.  72.     FACE  OF  Geomys  bursarius,  SHOWING  GROOVED  UPPER  INCISORS 

AND    OPENINGS    OF   CHEEK-POUCHES. 

FIG.  73.     FACE  OF  7"komomys  talpoides,  SHOWING  SMOOTH  UPPER  INCISORS 

AND    OPENINGS    OF   CHEEK-POUCHES. 

sub-region  of  the  Holarctic,  where  they  are  represented  by  a 
distinct  sub-genus  (Teonoma).  The  round-tailed  musk-rat  of 
Florida  (Neofiber)  is  an  exclusively  Sonoran  type,  although  it  is 
regarded  by  Dr  Merriam  merely  as  a  sub-genus  of  Microtus. 
Highly  characteristic  of  the  region  are  the  pouched  rats,  consti- 
tuting the  genera  GeomyidcR.  Of  these,  the  typical  genus  Geomys  * 
extends  northwards  into  the  Transition  zone  and  southwards  into 
Central  America ;  while  the  nearly-allied  Thomomys>  in  which  the 
upper  incisor  teeth  are  smooth  instead  of  grooved,  penetrates 
into  the  Canadian  sub-region  of  the  Holarctic,  although  unknown 

1  Subdivided  into  eight  genera  by  Merriam,  North  American  Fauna,  Part 
viii.,  Washington  (1895). 


X.]  ITS   FAUNA.  367 

in  Central  America.  Both  these  genera  are  represented  in  the 
Pliocene  of  the  Sonoran  area.  In  the  same  family  the  three 
genera  of  kangaroo-rats  known  as  Dipodomys,  Perodipus,  and 
Microdipodops  appear  to  be  confined  to  the  region ;  and  the  same 
is  the  case  with  the  allied  genus  Hderomys,  although  Perognathus 
passes  northwards  into  the  Transition  zone. 


FlG.    74.      UNDER  SURFACE   OF  LEFT   FORE-FOOT  OF    Geomys. 

In  the  Ungulata,  the  deer  belonging  to  the  peculiar  American 
genus  Cariacus  are  very  abundant  in  the  Sonoran  region  (where 
those  of  the  typical  genus  Cervus  are  entirely  wanting),  although 
they  also  range  into  the  Canadian  sub-region  of  the  Holarctic,  and 
extend  right  through  South  America.  Since,  however,  they  are 
wanting  in  the  earlier  Tertiary  deposits  of  the  latter  area,  as  they 
are  at  all  epochs  in  the  Old  World,  there  can  be  little  hesitation  in 
regarding  them  as  essentially  Sonoran  types.  Even  more  decidedly 
is  this  the  case  with  the  prongbuck  (Antilocapra),  the  sole  type  of 
the  family  Antilocapridce,  which  is  distinguished  from  the  Bovidce. 
by  the  horn-sheaths  of  the  males  being  branched  and  periodically 
shed  from  their  bony  supports.  Although  the  prongbuck  pene- 
trates a  considerable  distance  into  the  Canadian  sub-region  of  the 
Holarctic,  its  true  home  is  the  prairie-district  of  the  Sonoran  lying 


368 


THE   SONORAN    REGION. 


[CHAP. 


to  the  westward  of  the  Mississippi.  Possibly  a  small  deer-like 
animal  from  the  Tertiaries  of  the  same  area  known  as  Cosoryx, 
may  have  been  the  ancestral  stock  of  the  prongbuck.  Lastly,  the 
peccaries  (Dicotyles),  which  are  now  chiefly  South  American,  appear 
to  have  been  originally  Sonoran  types  which  have  migrated  south- 
wards ;  their  fossil  remains  being  common  in  the  Tertiaries  of  the 
United  States,  whereas  they  are  unknown  in  South  America  before 


FlG.   75.     HEAD  OF  MALE  MULE-DEER  (Cariacus  macrotis). 

the  Plistocene.  Their  near  affinity  to  the  earlier  Tertiary  pigs  of 
the  Old  World  indicates  that  at  a  more  remote  date  they  spread 
from  a  more  northerly  starting-point. 

With  regard  to  the  armadillo  (Tatusia)  found  in  the  Sonoran, 
this  is  clearly  a  very  recent  immigrant  from  the  Neogseic  realm  ; 
and  although  opossums  (Didelphys)  were  abundant  in  North 
America  during  the  early  portion  of  the  Tertiary  epoch,  it  is  not 
improbable  that  the  same  explanation  will  hold  good  for  their 
existing  Sonoran  representatives. 


X.] 


ITS  FAUNA. 


369 


The  following  list  includes  such  exclusively  New  World  genera 
of  mammals  (apart  from  bats)  which  are  represented  in  the 
Sonoran  area;  those  which  may  be  regarded  as  more  or  less 
nearly  confined  to  this  region  being  printed  in  italics.  To  appre- 
ciate fully  the  significance  of  this  list,  reference  must,  however,  be 
made  to  the  series  of  Holarctic  genera  given  on  p.  360,  which 


FIG.   76.     HEAD  OF  MALE  PRONGBUCK  (Antilocapra  americana). 

are  more  or  less  completely  restricted  to  the  Canadian  sub-region 
of  that  great  region,  and  the  intervening  Transition  zone.  The 
Sonoran  list  is  as  follows,  viz. : — 

Insectivora. 

SORICID^E. 

Notiosorex.     Also  Central  America. 
Blarina.     Enters  Canadian  sub-region  of  Holarctic. 
L.  24 


370  THE   SONORAN    REGION.  [CHAP. 

Insectivora  (cont.). 
TALPID^E. 

Scalops.     Enters  Transition  zone. 

Scapanus.     Enters  Canadian  sub-region  of  Holarctic. 

Carnivora. 

PROCYONID^E. 

Bassariscus.     Enters  Transition  and  Central  America. 
Procyon.     N.  to  S.  America. 
Nasua.     Also  South  American. 

MUSTELID^E. 

Spilogale.     Enters  Transition  and  Central  America. 

Conepatus.     Texas  to  Patagonia. 

Mephitis.       Extends    into    Canadian    sub-region    and 

Central  America. 
Taxidea.     Enters  Holarctic. 

Rodentia. 


Cynomys.     Extends  into  Holarctic. 

MURID^E. 

Rhithrodontomys. 

Sitomys.     The  whole  of  America. 

Sigmodon.     Southwards  to  Ecuador. 

Neotoma.     Ranges  into  Holarctic. 

Neofiber. 
GEOMYID^E. 

Geomys.      Extends   into   Transition   zone   and  Central 
America. 

Thomomys.     Ranges  into  Canadian  sub-region. 

Dipodomys. 

Perodipus. 

Microdipodops. 

Perognathus.     Ranges  into  Transition  zone. 

Heteromys. 

Ungulata. 

A  NTIL  OCA  PRIDJE. 

Antilocapra.     Ranges  into  Canadian  sub-region. 


X.]  EXTINCT    MAMMALS.  37! 

Ungulata  (cont.). 


Cariacus.     Greater  part  of  America. 

DlCOTYLID^E. 

Dicotyles.     Also  South  American. 
Edentata. 

DASYPODID^E. 

Tatusia.     South  American. 

Marsupialia. 

DlDELPHYID^E. 

Didelphys.     South  American. 

Although  the  Transition  zone  undoubtedly  forms  an  unsatis- 
factory item  in  regard  to  the  distinctness  of  the  Sonoran  region, 
yet  when  we  look  at  the  difference  of  its  mammalian  fauna  as  a 
whole  from  that  of  the  Canadian  sub-region  of  the  Holarctic,  and 
the  close  similarity  between  the  latter  and  the  fauna  of  northern 
Europe  and  Asia,  there  can  be  but  little  hesitation  in  regard  to 
the  acceptance  of  Dr  Merriam's  view  that  the  Sonoran  is  a  valid 
zoological  region  of  the  Arctogaeic  realm. 

In  a  previous  chapter  the  groups  of  mammals,  both  living  and 
extinct,  confined   to   the   eastern   division   of  the 
Arctogaeic  realm  have  been  already  noticed,  while     oraupa  of 
in  the  present  one  reference  has  been  made  to  such 


existing  types  as  are  restricted  to  the  western  half    of  Western 

,     6     /F  .  .,  Arctogaea. 

of  the  same  realm.  It  now  remains  to  consider 
briefly  some  of  the  leading  extinct  groups  which  are  found  only  in 
the  latter  area  ;  and  the  consideration  of  these  comes  most  appro- 
priately here,  seeing  that  the  majority  of  these  peculiarly  American 
types  are  of  Sonoran  origin,  a  large  number  of  their  remains 
having  been  obtained  from  the  States  of  New  Mexico,  Kansas, 
Nebraska,  and  Dakota,  which  lie  within  that  region,  or  from 
Colorado,  Wyoming,  and  Montana,  which  are  situated  within  the 
Transition  zone. 

Although,  in  common  with  the  higher  Primates,  lemuroids  are 
now  quite  unknown  in  North  America,  they  were  well  represented 
there  during  the  Puerco  epoch  of  the  lower  Eocene  by  three 
families.  The  first  of  these  —  the  Chriacida  —  includes  animals 

24—2 


372 


THE   SONORAN    REGION. 


[CHAP. 


X.]  EXTINCT   MAMMALS.  373 

having  the  same  number  of  teeth  as  the  allied  Tertiary  European 
family  Adapidce,  but  all  characterised  by  their  more  primitive 
structural  features.  Indeed  these  early  lemuroids  appear  to 
present  considerable  resemblances  to  the  creodont  Carnivora, 
and  differ  from  all  the  other  members  of  the  sub-order  to  which 
they  belong  by  the  great  elongation  of  the  bony  symphysis  con- 
necting the  two  branches  of  the  lower  jaw  at  the  chin.  Several 
other  genera,  in  addition  to  the  typical  Chrtacis,  are  assigned 
to  this  family.  The  second  group  is  that  of  the  Anaptomorphidce, 
which  is  represented  in  the  Puerco  Eocene  by  a  genus  known  as 
Indrodon,  and  in  somewhat  higher  beds  by  the  typical  Anapto- 
morphus.  Although  in  other  respects  coming  closer  to  existing 
types  than  is  the  case  with  the  Chriacidcz,  the  present  family  is 
broadly  distinguished  by  the  tritubercular  structure  of  the  upper 
molar  teeth.  The  third  peculiar  North  American  family  of  the 
lemuroids  is  that  of  the  Mixodectidcz,  typically  represented  by 
Mixodectes  of  the  Puerco  Eocene. 

Among  the  extinct  creodont  Carnivora  there  are  two  families 
apparently  restricted  to  the  Tertiaries  of  North  America,  namely, 
the  Miacidce,  and  the  Mesonychidcz,  the  former  of  which  presents 
such  strongly  marked  affinities  to  the  modern  Carnivora  that  it 
is  frequently  assigned  to  that  group.  The  second  family,  on  the 
other  hand,  as  represented  typically  by  the  genus  Mesonyx  of  the 
Uinta  or  lowest  Oligocene,  is  characterised  by  the  very  simple 
structure  of  the  whole  series  of  cheek-teeth,  which  are  not  unlike 
the  pre-molars  of  some  of  the  higher  carnivores.  One  of  the 
species  of  the  typical  genus  attained  dimensions  as  large  as  those 
of  a  bear.  In  the  widely  distributed  family  Hyanodontida,  an 
exclusively  North  American  genus  is  Patriofelis,  which  is  regarded 
as  a  specialised  offshoot  from  Oxyana. 

Among  the  ungulates  there  are  several  extinct  families  con- 
fined to  North  America.  In  the  group  forming  a  transition 
between  the  pigs  and  the  ruminants  there  is  first  of  all  the  family 
of  the  oreodonts  (Cotylopida),  which  make  their  appearance  in 
the  middle  Oligocene,  and  continue  to  the  Miocene  and  lower 
Pliocene1.  These  ungulates,  which  were  allied  to  the  genus 

1  By  American  geologists  the  term  Oligocene  is  not  generally  used,  so  that 
the  whole  of  the  Tertiary  strata  are  classed  as  Plistocene,  Pliocene,  Miocene, 


374 


THE    SONORAN    REGION. 


[CHAP. 


1 


X.]  EXTINCT   MAMMALS.  375 

Ancodus,  common  to  the  Tertiaries  of  both  hemispheres1,  and 
were  represented  by  a  large  number  of  generic  types,  have 
crescentic  columns  to  the  short-crowned  cheek-teeth,  the  upper 
molars  usually  carrying  four  such  columns  ;  while  the  lower  canine 
is  approximated  to  the  incisors,  its  usual  form  and  function  being 
assumed  by  the  first  pre-molar.  The  last  upper  pre-molar  is 
simpler  than  the  molars ;  and  while  the  feet  have  usually  four  toes 
each,  in  the  typical  genus  Cotylops  a  rudiment  of  the  thumb  is 
retained  in  the  front  pair,  as  in  Ancodus.  In  Cotylops  and  most 
of  the  other  genera  the  molars  of  the  upper  jaw  have  but  four 
columns,  but  in  Protoreodon  there  are  five ; — a  feature  serving  to 
connect  the  family  with  the  Anthracotheriida,  from  which  group 
the  oreodonts  are  probably  descended.  A  nearly-allied  but  more 
specialised  family  is  that  of  the  Agriochceridce,  as  represented  by 
the  genus  Agriochcerus* ,  of  the  upper  and  middle  Oligocene,  in 
which  the  toes  were  developed  into  claws,  instead  of  being 
incased  in  hoofs.  Here  it  may  be  mentioned  that  while  the 
peccaries  (Dicotylida)  are  now  exclusively  New  World  types,  and 
the  pigs  (Sutdce)  restricted  to  the  Old  World,  the  Tertiaries  of  both 

and  Eocene.  Introducing  the  former  term,  the  series  may  be  approximately 
classified  as  follows,  viz. : 

PLISTOCENE.  Equus  Beds.     Eqtius^  Elephas  primigenius. 

PLIOCENE  (Upper).  Blanco  Series.     Pliauchenia. 

(Lower).  Loup-Fork.     Protohippus,  Hipparion. 

MIOCENE.  Deep  River.    Anchitherium^  First  Mastodons. 

OLIGOCENE          (Upper).  John  Day.     Miohippus,  Ancodus. 

(Middle).  White  River.     Agrwchcerus,  Titanotherium. 

(Lower).  Uinta.     Amynodon,  Mesonyx. 
EOCENE           (Up.  and  Mid.).     Bridger.     Pachynolophus,  Paltzosyops. 

(Lower).  Wahsatch.     Hyracotheritim,  Coryphodon. 

(Lowest).  Puerco.     Neoplagiaulax ,  Poly  mastodon. 

In  this  series  the  Deep  River  beds  are  identified  with  the  European 
Miocene  (supra^  p.  117)  by  the  presence  of  Anchitheritim,  and  the  first  appear- 
ance of  Mastodon  ;  while  the  existence  of  Ancodus  in  the  John  Day  and  Upper 
White  River  beds  correlates  them  with  the  Upper  and  Middle  Oligocene, 
Hyracotherium  and  Coryphodon  serving  to  identify  the  Wahsatch  beds  with  the 
Lower  Eocene. 

1  Supra,  p.  161. 

2  Equal  Artionyx. 


376 


THE   SONORAN    REGION. 


[CHAP. 


hemispheres  contain  intermediate  types  such  as  Hyotherium  and 
Chcerohyus  which  are  apparently  the  ancestral  stock  of  both 
families. 


FIG.  79.     FRONT  VIEW  OF  RIGHT  HIND  FOOT  OF  Agriochcerus. 

Another  very  peculiar  type  of  North  American  Tertiary  ungu- 
lates is  represented  by  Protoceras,  from  the  upper  division  of  the 
White  River  Oligocene,  which  forms  a  family  (Protoceratidce}  by 
itself.  In  these  creatures  the  feet  approximate  to  the  ruminant 
type;  but  the  skull,  as  shown  in  the  accompanying  illustration, 


FIG.  80.     SKULL  OF  Protoceras,  WITHOUT  THE  LOWER  JAW. 

has  at  least  two  pairs  of  large  bony  processes,  probably  covered  in 
life  with  horns,  and  a  pair  of  large  upper  tusks,  in  both  of  which 
respects  it  exhibits  a  curious  parallelism  with  the  perissodactyle 


X.]  EXTINCT    MAMMALS.  377 

ungulates.  No  trace  of  these  singular  artiodactyles  has  hitherto 
been  detected  in  the  Old  World. 

The  Camelidce.  seem  to  have  been  primarily  a  North  American 
family,  which  originated  in  the  Sonoran  region,  and  of  which  one 
branch  (Lama)  subsequently  migrated  south,  while  the  other 
(Camelus)  crossed  Bering  Strait  into  the  Old  World.  In  the 
upper  Pliocene  there  occurs  Pliauchenia,  with  only  three  lower 
pre-molars,  and  in  the  lower  Pliocene  Loup-Fork  beds  Procamelus 
with  four  of  these  teeth  ;  while  the  earliest  representative  of  the 
family  is  Leptotragulus  of  the  Uinta  Oligocene. 

In  the  perissodactyle  section  of  the  same  order  the  family 
Titanotheriidce  is  mainly  North  American,  although,  as  stated  on 
page  107,  a  representative  of  the  typical  genus  Titanotherium  has 
been  discovered  in  the  Tertiaries  of  the  Balkans.  Titanotherium 
includes  huge  rhinoceros-like  animals,  with  low-crowned  molar 


FIG.  8r.     RIGHT  UPPER  MOLAR  TOOTH  OF  Palaosyops, 

teeth  of  the  type  of  those  of  Chalicotherium,  and  frequently  having 
the  nasal  region  of  the  skull  surmounted  by  large  bony  protuber- 
ances. The  genus  is  characteristic  of  the  Uinta  and  the  lower 
division  of  the  White  River  Oligocene.  An  earlier  type  of  the 
same  family  is  typified  by  the  smaller  and  less  specialised  hornless 
animals  from  the  Bridger  Eocene,  known  as  Palaosyops,  which, 
together  with  certain  allied  forms,  constitute  a  peculiar  sub-family 
confined  to  America.  This  family,  like  the  camels,  appears  there- 
fore to  have  originated  in  the  Sonoran  region,  whence  a  few 
representatives  wandered  eastwards  into  the  Old  World. 

In  the  generalised  ungulate  sub-order  termed  Amblypoda, 
of  which  the  lower  Eocene  coryphodons  were  the  earliest 
representatives,  North  America  possesses  an  absolutely  peculiar 


THE   SONORAN    REGION.  [CHAP. 

family  in  the  UintatheriidcR.  These  were  huge,  somewhat  ele- 
phantine ungulates,  with  five  short  toes  to  each  foot,  long  tusk- 
like  canine-teeth  in  the  upper  jaw,  and  the  skull  surmounted  with 
three  pairs  of  large  bony  protuberances ;  their  molar  teeth  being  a 
specialised  form  of  the  Coryphodon  type.  They  occur  in  the 
middle  division  of  the  Bridger,  or  the  one  above  the  zone  yielding 
Coryphodon,  and  may  accordingly  have  been  the  descendants  of 
that  genus.  These  uintatheres  appear  to  have  been  restricted  to 
the  "  Bad  Lands  "  of  the  Sonoran  region  and  adjacent  districts  of 
the  Transition  zone. 


FlG.    82.       EXTREMITY   OF    SKULL  OF   UintCltherium ,  TO   SHOW  UPPER   TUSKS. 

Passing  by  certain  other  forms  of  less  interest,  attention  may 
be  directed  to  a  peculiar  group  of  aberrant  mammals  forming 
the  Tillodontia,  which  appear  to  be  mainly  North  American,  and  of 
which  the  serial  position  cannot  be  precisely  determined.  They 
are  restricted  to  the  Eocene,  and  seem  to  combine  the  characters 
of  the  modern  Ungulata,  Rodentia,  and  Carnivora.  In  the  genus 
Anchippodus,  forming  the  type  of  the  family  AnchippodontidcB,  the 
skull  approximates  to  that  of  a  bear,  the  cheek-teeth  are  of  an 
ungulate  type,  and  there  is  a  pair  of  large  chisel-like  incisors 
(preceded  by  a  small  functionless  upper  pair)  in  each  jaw,  very 
similar  to  those  of  the  rodents  and  hyraces.  A  second  family, 


X.I  LIST   OF   THE    FAUNA.  379 

Psittacotheriida,  is  represented  by  the  genera  Psittacotherium  and 
Calamodon,  in  which  the  cheek-teeth  grew  permanently,  instead  of 
developing  roots. 

The  following  list  exhibits  the  chief  families  or  minor  groups 
of  characteristically  North  American  mammals,  which  are  either 
entirely  wanting  or  but  sparingly  represented  in  the  Old  World ; 
those  that  are  extinct  being  indicated  by  a  t,  and  the  absolutely 
characteristic  forms  being  printed  in  italics. 

Primates. 

t  Anaptomorphidce.     Anaptomorphus. 
t  Mixodectida.     Mixodectes. 
t  Chriacidce.     Chriads. 

Carnivora. 

Procyonidae.      Represented   in   the   Old   World   only   by 

^Elurus. 

t  Miaridce.     Miacis,  Didynrictis. 
t  Mesonychidcz.      Mesonyx. 
fHyaenodontidse.     Patriofelis. 

Rodentia. 

Haplodon  tidce. 
Geomyidce. 

Ungulata. 

Dicotylidce.     Represented  by  ancestral  types  in  Tertiaries 

of  E.  Hemisphere. 

t  Cotylopida.     Cotylops,   Mesoreodon,   Protoreodon. 
t  Agriochceridce.     Agrioch&rus. 
t  Protoceratida.     Protoceras. 

Camelidae.     t  Pliauchenia,   \Procamelus,   M^eptotraguius. 

Antilocaprida.     Antilocapra,  t  Cosoryx. 
t  Titanotheriidae.     Palceosyops,  Limnohyops,  Telmatotherium. 
t  Uintatheriidce.      Uintatherium. 

Tillodontia. 

t  Psittacotheriida.     Psittacotherium,    Calamodon. 
t  A  nchippodontidce.     A  nchippodus. 


380  THE   SONORAN    REGION.  [CHAP. 

In  an  earlier  chapter1  a  list  has  been  given  of  the  leading 
Distinctness  mammalian  families  common  to  the  two  divisions 
of  the  Region.  of  Arctogaea,  and  since  in  the  foregoing  chapter  it 
has  been  shown  that  many  of  the  peculiar  American  families 
are  more  or  less  intimately  related  to  some  of  those  common  to 
the  two  areas,  it  is  manifest  that  throughout  the  Tertiary  period 
eastern  and  western  Arctogaea  must  have  had  a  land-connection 
towards  the  north,  so  that  there  was  an  interchange  of  the  fauna 
of  the  more  northern  districts.  Those  American  types  which 
penetrated  as  far  south  as  what  is  now  the  Sonoran  area  would, 
however,  naturally  tend  to  become  isolated,  and  thus  develop 
into  the  families  which  may  be  regarded  as  characteristic  of  that 
region.  So  far,  therefore,  from  this  area  being  merely  a  part  of  a 
so-called  Nearctic  region,  there  are  indications  that  it  was  differ- 
entiated from  the  Holarctic  at  a  time  when  the  existing  zoological 
regions  of  the  eastern  half  of  the  Arctogaeic  realm  were  still  unde- 
fined. Indeed  from  the  community  of  the  Pliocene  fauna  of 
southern  Europe,  Asia  Minor,  Persia,  northern  India,  and  south 
China,  it  seems  probable  that  the  only  divisions  of  the  Arctogaeic 
realm  that  could  have  been  attempted  would  have  been  into  (i)  a 
Sonoran  region,  (2)  a  Holarctic  region,  comprising  the  northern 
districts  of  America,  Asia,  and  Europe,  (3)  what  may  be  termed  a 
Mediterraneo-Oriental  region,  including  southern  Europe,  north 
Africa,  and  the  whole  of  southern  Asia ;  and  (4)  a  Malagasy 
region,  which  would  then,  or  perhaps  somewhat  earlier,  have 
included  Ethiopian  Africa. 

As   to  the  amount   of  interchange  which  took  place  during 
Dual  Origin      Tertiary  times  between  the  mammals  of  the  eastern 
of  Groups.  and  western  divisions  of  Arctogaea,  and  as  to  whether 

similar  generic  types  may  have  been  developed  independently  in 
the  two  areas,  it  is  almost  impossible  to  arrive  at  any  satisfactory 
conclusion.  The  suggestion  that  Equus  has  thus  been  independ- 
ently evolved  in  the  two  areas,  has  been  already  mentioned2,  and 
this  idea  receives  support  from  some  very  remarkable  observations 
recently  made  on  the  invertebrates  inhabiting  certain  European 
and  North  American  caves. 

1  Sttpra,  p.  176.  2  Supra,  p.  168. 


X.]  DUAL  ORIGIN    OF   GROUPS.  381 

With  a  quotation  from  Mr  G.  H.  Carpenter's  interesting 
paper1  on  this  subject,  the  present  volume  may  be  fitly  closed. 
After  describing  the  inhabitants  of  the  Mitchelstown  Cave,  in 
Ireland,  the  author  writes  that  the  spring-tail  (Lipurd]  "is  hardly 
to  be  separated  from  a  species  found  in  the  caves  of  Carniola,  and 
the  Sinella  is  almost  identical  with  one  inhabiting  the  caves  of 
North' America  ;  while  the  spider  is  apparently  the  same  as  a  cave- 
dweller  from  the  Mediterranean  district  of  southern  France,  which 
probably  occurs  in  the  North  American  caverns  also.  Had  we  to 
do  with  animals  of  the  upper  fauna,  these  results,  though  highly 
interesting,  would  not  be  without  parallel  in  species  already 
known....  But  the  occurrence  of  cave-dwelling  species  with  so 
wide  a  range  is  a  truly  remarkable  phenomenon.  The  caves 
cannot  be  of  any  great  geological  age.  Any  possible  geographical 
connection  which  would  permit  the  migration  of  subterranean 
animals  between  southern  Europe  and  Ireland,  or  between  Ireland 
and  North  America,  seems  altogether  out  of  the  question  within 
any  period  during  which  the  fauna  can  have  been  specifically 
identical  with  that  of  the  present  day.  The  only  conclusion  is 
that  from  ancestors,  presumably  of  the  same  genus,  which  took  to 
an  underground  life  in  such  widely-separated  localities,  the  similar 
conditions  of  the  caves  have  evolved  descendants  so  similar  that, 
when  compared,  they  cannot  or  can  hardly  be  specifically  distin- 
guished from  each  other.  Should  the  identifications  stand  the 
test  of  a  comparison  of  types,  we  shall  have  proof  that  the  inde- 
pendent development  of  the  same  species,  under  similar  conditions, 
but  in  widely-distant  localities,  has  taken  place.  It  must  be 
granted,  however,  that  cave-conditions  are  so  marked  and  excep- 
tional, that  it  might  not  be  safe  to  argue  from  them  as  to  what 
may  have  occurred  in  the  upper  world." 

Although  the  author  of  this  passage  is  perfectly  correct  in  his 
statement  that  there  is  a  vast  difference  between  cave-life  and 
open-air  life,  yet  if  animals  which  appear  to  belong  to  one  and  the 
same  species  can  be  proved  to  have  had  a  dual  origin  in  the  one 
case,  it  can  scarcely  be  considered  impossible  that  similar  instances 
may  occur  in  the  other.  And  if  such  dual  origins  exist  among 

1  Irish  Naturalist,  Vol.  IV.  pp.  25 — 35  (1895). 


382  THE   SONORAN    REGION.  [CHAP.  X. 

species,  there  is  surely  no  reason  why  they  should  not  occasionally 
occur  in  the  case  of  genera.  It  would,  therefore,  seem  by  no 
means  improbable  that  the  species  of  the  genus  Equus  which 
inhabited  the  eastern  and  western  halves  of  the  northern  hemi- 
sphere during  the  close  of  the  Tertiary  period  may  have  been 
evolved  from  the  closely-allied  but  separate  ancestral  equine 
stocks. 

The  matter  does  not,  however,  by  any  means  end  here.  In 
an  earlier  chapter1  it  has  been  shown  that  the  same  species  of  a 
genus  of  fish  (Galaxias)  occurs  in  countries  so  remote  from  one 
another  as  New  Zealand  and  Australia  on  the  one  hand,  and 
Patagonia  on  the  other.  With  the  evidence  of  the  cave-animals 
before  us,  is  it  absolutely  impossible  that  these  apparently  iden- 
tical fishes  can  have  been  evolved  independently  of  each  other  ? 
Should  this  be  so,  it  will  engender  increased  caution  in  drawing 
any  inferences  as  to  former  land-connection  from  the  evidence 
of  single  animals.  But  such  instances  of  independent  evolution, 
if  they  do  occur,  must  be  of  extreme  rarity,  and  will  in  no  case 
interfere  with  deductions  drawn  from  the  presence  of  a  number 
of  allied  species  or  genera  of  animals  in  distant  countries. 

1  Suprh,  p.  125.     Recently  another  species  has  been  described  from  South 
Africa.     See  Steindachner,  SB.  Ac.  Wien,  vol.  ciii.  p.  460  (1894). 


APPENDIX. 


LIST  OF  WORKS  AND  PAPERS  MOST  FREQUENTLY  REFERRED  TO 
IN  THIS  VOLUME. 

1.  ALLEN,  J.  A.    The  Geographical  Distribution  of  Mammals.  Bull. 

U.  S.  Geol.  Survey,  vol.  IV.  nos.  2  and  4,  pp.  313—376,  (1878). 

2.     The    Geographical     Distribution    of    North    American 

Mammals.     Bull.  Amer.  Mus.,  vol.  IV.  pp.  199—243,  (1892). 

3.  ANONYMOUS.     Antarctica;  a  Supposed  Former  Southern  Conti- 

nent.    Natural  Science,  vol.  III.,  pp.  54—57,  (1893). 

4.     The  Nearctic  Region  and  its  Mammals.     T.  c.  pp.  288 — 

292. 

5.  BEDDARD,  F.  E.     A  Text-Book  of  Zoogeography.     (Cambridge 

Natural  Science  Manuals}.     Cambridge,  1895. 

6.  BLANFORD,  W.   T.    The  African    Element    in    the    Fauna  of 

India.     Ann.  Mag.  Nat.  Hist.  ser.  4,  vol.  XVIII.,  pp.  277 — 294, 
(1876). 

7.     Note  on  the  "Africa- 1 ndien"  of  A.  von  Pelzeln,  and  on 

the  Mammalian  Fauna  of  Tibet.     Proc.  Zool.  Soc.   1876,  pp. 
631—634. 

8.     Anniversary  Address  to  the  Geological  Society.     Proc. 

Geol.  Soc.  1890,  pp.  43—110. 

9.     The  Age  of  the  Himalayas.     Geol.  Mag.  decade  3,  vol.  IX., 

pp.  161 — 168,  (1892). 

10.  — On   a   Stag  (Cervus  thoroldi}  from  Tibet,  and   on   the 

Mammals  of  the  Tibetan  Plateau.     Proc.  Zool.  Soc.  1893,  pp. 
444-449. 

11.  BOURNS,  F.  S.,  and  DEAN,  C.  W.     Preliminary  Notes  on  Birds 

and  Mammals  collected  in  the  Philippine  Islands.     Occasional 
Papers  Minnesota  Acad.  vol.  I.,  pp.  I  —64,  (1894). 

12.  BULMAN,  G.  W.     The  Effect  of  the  Glacial  Period  on  the  Fauna 

and  Flora  of  the  British  Islands.     Natural  Science,  vol.   ill., 
pp.  261 — 266,  (1893). 


384  APPENDIX. 


13.  CARPENTER,   G.    H.     Nearctic   or   Sonoran  ?     Natural  Science, 

vol.  v.,  pp.  53—57,  (1894). 

14.  EVERETT,  A.  H.     A  Nominal  List  of  the  Mammals  inhabiting 

the   Bornean    Group   of  Islands.     Proc.  Zool.  Soc.    1893,   pp. 
492—496. 

15.  FORBES,  H.  O.     The    Chatham    Islands;    their    Relation   to    a 

Former  Southern  Continent.     Supplemental  Papers  R.  Geogr. 
Soc,  1893,  pp.  607—637. 

1 6.  HEDLEY,    C.     On   the    Relation   of    the    Fauna  and    Flora    of 

Australia  to  those  of  New  Zealand.     Natural  Science,  vol.  III., 
pp.  187— 191,  (1893). 

17.  HEILPRIN,  A.     The  Geographical  and  Geological  Distribution 

of  Animals.     International  Scientific  Series,  London,  1887. 

1 8.  HUXLEY,   T.    H.     On   the    Classification    and    Distribution    of 

Alectoromorphas  and   Heteromorphas.     Proc.  ZooL   Soc.   1868, 
pp.  294—319. 

19.  MERRIAM,  C.  H.     The  Geographic  Distribution  of  Life  in  North 

America,    with    special    reference    to    the    Mammalia.     Proc. 
Biol.  Soc.  Washington,  vol.  VI I.,  pp.  i — 64,  (1892). 

20.     Laws  of  Temperature  Control  of  the  Geographic  Dis- 
tribution of  Terrestrial  Animals  and  Plants.     Nat.  Geogr.  Mag. 
vol.  VI.,  pp.  229 — 238,  (1894). 

21.  NEHRING,  A.     Ueber  Saugethiere  von  den  Philippinen,  nament- 

lich  von   der  Palawan-Gruppe.     Sitzber.   Ges.  Naturf.  Berlin, 
1894,  pp.  I79—I93- 

22.  OGILBY,   J.    D.     Catalogue   of  Australian    Mammals.     Sydney, 

1894. 

23.  OSBORN,  H.  F.     The  Rise  of  the  Mammalia  in  North  America. 

Stud.  Biol.  Lab  or  at.  Columbia  College,  Zoology,  vol.  I.,  art.  2, 

(1893). 

24.     ,  and  EARLE,  C.     Fossil  Mammals  of  the  Puerco  Beds. 

Bull.  Amer.  Mus.  vol.  VIL,  pp.  i — 70,  (1895). 

25.  SCHARFF,  R.  F.     On  the  Origin  of  the  Irish  Land  and  Fresh- 

water Fauna.     Proc.  Irish  Acad.  ser.  3,  vol.  III.,  pp.  479 — 485, 
(1894). 

26.  SCLATER,  P.  L.     On  the  General  Geographic  Distribution  of  the 

Members  of  the  Class  Aves.    Journ.  Linn.  Soc.  Zool.  vol.  II., 
pp.  130—145,  (1858). 

27.     The    Geographical    Distribution    of    Mammals.     Man- 
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28.     W.    L.     The    Geography    of    Mammals.     Geographical 

Journal,  1894,  1895. 


APPENDIX.  385 


29.  SHARPE,  R.  B.     On  the  Zoo-Geographical  Areas  of  the  World, 

illustrating  the  Distribution  of  Birds.    Natural  Science,\o\.  III., 
pp.  100—108,  (1893). 

30.  THOMAS,  O.  T.     The  Mammals  of  the  Solomon  Islands.    Proc. 

Zool.  Soc.  1888,  pp.  470—484. 

31.     Preliminary  Diagnosis  of  New  Mammals  from  Northern 

Luzon,  collected  by  Mr  J.  Whitehead.     Ann,  Mag.  Nat.  Hist. 
ser.  6,  vol.  XV.  pp.  160 — 164,  (1895). 

32.  WALLACE,  A.   R.     The  Geographical  Distribution  of  Animals. 

London,  1876,  2  vols.  8vo. 

33.     Island  Life.     London,  1880,  8vo. 

34.     What    are    Zoological    Regions?      Nature,    vol.   XLIX., 

pp.  6 10 — 613. 

35.     The  Palaearctic  and  Nearctic  Regions  compared  as  re- 
gards the  Families  and  Genera  of  their  Mammalia  and  Birds. 
Natural  Science,  vol.  IV.,  pp.  433 — 445,  (1894). 

36.  ZITTEL,  K.  A.  VON.   Die  geologische  Entwickelung,  Herkunft,  und 

Verbreitung  der  Saugethiere.     Sitzber.  bayer.  Akad.  vol.  xxill., 
pp.  137—198,  (1893). 


L.  25 


INDEX. 


Aard-varks,  187,  249,  256,  268 

Aard-wolf,  235 

Abderites,  in 

Abderitidcz,  1 1 1 

Acaremys,  90 

Acdestis,  no 

Acomys,  198 

Ac  other  uhim.)  192 

Acrobates,  37 

Adapis,  181,  191,  219 

Addax,  245,  324,  358 

Adelphomys,  91 

^  157,  191,  202 
$)  321 
212,  275 
sEpyceros,  245 
sEpyornis,  222 
sEpyprymnus,  36 
Age  of  animal  groups,  7 
Agriochceridiz,  375 
Agriochceriis,  375 
Agutis,  89,  137 
Alactaga,  323,  351 
^&w,  164,  315 
Amblotherium,  53 
Amblypoda,  173,  377 
Amblyrhiza,  87,  137 
Ameghino,  on  age  of  Santa  Cruz  fauna, 

68 

Ammodorcas,  245 
AmphictiSy  191 
Amphicyon,  158,  191 
.4  ?«/>//  icy  nod  on  ,193 
Amphidozotherium,  156,  191 
Amphilestes,  53 
Amphiproviverra,  109 
Amphisbcend)  132 
Amphisbcenidtz,   distribution  of,    132; 

fossil,  132 
AmphitheriidcB,  52 
Amphitherium,  53 


itragulus,  1 94 

^?  nalcitheriu  m,  105 

Anaptomorphidce,  373 

Anaptomorphus,  373 

Anchilophus,  166,  192,  193 

Anchippodontid&i  378 

AnchippodttS)  378 

Anchithei'ium,  166,  196 

Ancodus,  161,  193,  203,  375 

Ancyclotherium,  172,  200 

Anoa,  47,  164 

Anomaluridce,  237 

Anomalurus,  238 

Anomodontia,  151 

AnoplotheriidcZ)  185 

Anoplotherium,  185,  193 

Anops,  132 

Antarctica,  56,  128;  extent  of,  129 

Antarctogsea,  26 

Anteaters,  101,  136 

^4  ntech  inom  ys,  3  9 

Anthracotheriida,  161,  192 

Anthracotherium,)  163,  192,  193,  194,. 

203 

Anthropopithecus,  180,  202,  231,  256 
Antillia,  139 

Antillean  sub-region,  136,  137 
Antilocapra,  367 
Antilocapridce,  367 
Antelopes,  African,  242 
Amirosorex,  272 
Africa   connected   with    S.   America, 

127 

Africa,  route  of  migration  into,  256 
Aplin,     Mr,     on    mammals    crossing 

rivers,  14 

Aquilonian  region,  26 
Arabia,  South-eastern,  fauna  of,  262 
Arapaima,  134 
Arch&lurus,  157 
Archceoniys,  91 


INDEX. 


387 


Arctictis,  273 

Arctic  sub-region,  360 

Arctic  zone,  310 

Arctocyon,  159 

Arctogale,  273 

Arctogsea,  25,  26,  27,  144,  210 

Arctogsea,  Eastern,  179 

Arctogasan  region,  26 

Arctogaeic  fauna,  features  of,  147 

Arctogseic  realm,  27,  144 

Arctotherium,  72 

Arctomys,  159,  267,  351 

Arc t onyx,  276 

Area,  2 

Armadillo,  giant,  136 

Armadillos,  94,  368 

Artionyx,  375 

Arui,  35« 

Arvicanthis,  239 

Asses,  359 

Astrapotheria,  81 

Astrapotheriutn,  81 

Atherura,  278 

Atlantic,  its  recent  origin,  24 

Atlantosaurus,  35,  151 

Auk,  347 

Anlacodus,  91,  240 

Australia,  how  it  received  its  fauna, 
54;  its  connection  with  S.  America, 
55»  125;  its  relations  with  New 
Zealand  and  other  islands,  58,  59 

Australian  region,  25,  27,  31 

Austro-Columbian  region,  25 

Austro-Malayan  region,  27,  28,  45 

Avahi,  217 

Awantibo,  256 

Aye-aye,  219 

Babirusa,  47 

Bachitherium,  164,  192 

Badgers,  321 

Balanoptera,  68 

Banded  anteater,  40 

Bandicoots,  38 

Banteng,  278 

Barbets,  254 

Barriers  to  dispersal,  10,  14 

Bassariscus,  73,  136,  364 

Bathyergus,  23  r,  239 

B  atomy  s,  278 

Bats,  Notogseic,  42 ;  Neogseic,  70 

Baur,  Dr,  on  relations  of  Galapagos 

islands,  141 
Bdeogale,  235 
Bears,  American,  318;  their  absence 

from  Ethiopian  Africa,  258 
Beavers,  313,  318 


Beddard,  on  earth-worms,  126,  311; 
on  Fernando  Noronha,  140 

Bering  Strait  indicates  a  line  of  con- 
nection between  Asia  and  America, 

178?  337 
Bettongia,  30 
Birds,    Malagasy,    254;     Ethiopian, 

254 

Bison,  314 

Black-buck,  278 

Blanford,  Dr,  on  permanency  of  conti- 
nents, 24  ;  on  zoological  regions,  27  ; 
on  a  former  connection  between  the 
southern  continents,  128;  on  con- 
nection between  Africa  and  S. 
America,  128;  on  survival  of  old 
forms  in  Madagascar,  131;  on  the 
birds  of  Madagascar  and  Africa, 
254 ;  on  Oriental  sub-regions,  266 ; 
on  the  fauna  of  India,  267 ;  on  land 
connection  between  Ceylon  and 
Malaysia,  293 ;  on  the  Tibetan 
fauna,  354 

Blarina,  364 

Blastomeryx,  74 

Blesbok,  244 

Boa,  132 

Boas,  distribution  of,  132 

Bontebok,  244 

Boreal  region,  146,  309,  360;  zone, 
360 

Borhycena,  109 

Borneo,  its  fauna,  294 

Bos,  165,  206,  278,  314;  depressicornis, 
47,  164;  mindorensis,  47 

Boselaphus,  206 

Both  riospondyhis,  152 

Bothremys,  131 

Bothriceps,  152 

Bourbon,  213 

Bovidce,  164,  186,  196;  Holarctic,  314; 
Oriental,  279 

Brachydiastematotherium,  \  70 

Brachyurns,  \  36 

Brachyteles,  136 

Bradypodida,  100 

Brady  pus,  100 

Bramatherium ,  186,  204 

Brazilian  sub-region,  135 

British  Isles,  their  connection  with 
the  Continent,  339 

Brontornis,  60 

Bubalis,  206,  244,  358 

Budorcas,  324 

Buffalo,  279 

Bulman,  Mr,  on  the  submergence  of 
Britain,  350 

25—2 


388 


INDEX. 


Buphaga,  254 

Burmese  sub-region,  267 

Buried  river-channels  of  Britain,  339 

Cadurcotherium^  82,  170,  192 

Ccenolestes,  no 

C&notheriidce,  163,  192 

Ccenotherium,  163,  192,  193 

Calamodon,  379 

Callithrix,  136 

Caloprymnus,  36 

CamelidcE,  163,  185,  326,  377;  Neo- 

gseic,  74 

Camelus,  163,  185,  204,  377 
Camels,  326 
Campos  of  Brazil,  65 
Canadian  sub-region,  360 
Cane-rat,  91 

Canidce,  Oriental,  274;  Neogaeic,  72 
Cam's,    195,   202,    235,   274;   antarc- 

ticus,  140;  dingo,  ^i;frocyonides, 

355 

Cannabateomys,  91 

Capra,  206,  230,  280,  324,  359,  360 

Capreolus,  352 

Capivara,  88 

Capromyida,  90 

Capromys,  91,  137 

Caracal,  249 

Cardiotherium,  89 

Carcttochelyidcz,  62 

Carettochelys,  62 

Cariacus,  74,  344,  367 

Carioderma,  96 

Carnivora,  Arctogseic,  157;  Holarctic, 
312,  321;  Neogaeic,  71;  North 
American,  341 ;  Sonoran,  364 

Carpenter,  Mr,  on  cave-faunas,  381 

Carpincho,  88,  136 

Carpotnys,  278 

Carterodon,  91 

Castor,  313 

Castorida,  159,  313 

Castoroides,  87,  138 

Castoroidida,  87,  138 

Cassowaries,  48,  58 

Casuarius,  48 

Catamarca  beds,  67 

Catonyx,  105 

Caves  of  Lagoa  Santa,  66 

Ctf^a,  88 

Cavies,  88 

Caviidcz,  88 

Cave-faunas,  381 

Caxomistle,  136 

Cebidce,  69 

Cebochczrus,  192 


Celebes,  its  fauna,  47;  an  anomalous 

island,  49 
Cewas,  280 
Centetes,  220 
Centetidce,  131,  220 
Central  American  sub-region,  136 
Central  Asian  sub-region,  351 
Centres  of  evolution,  2  r 
Cephalogale,  191 
Cephalophus,  244 
Ceratodns,  133 
Ceratophora,  120 
Cercocebus,  231 
Cercoleptes,  72 
Cercomys,  91 
Cercopithecida,  231,  269;  distribution 

of,  180 

Cercopithecus,  231 
Cernaysian  Fauna,  153 
Cervicapra,  245 
Cervicaprince,  244,  245 
Cervidce,  164;  absence  of  in  Ethiopian 

region,    230,   258;  Holarctic,  315 ; 

Oriental,  280 
Cervulus,  198,  281 
CVrzw,  164,  280,  326,  335'.358>  359> 

367;  eustephanus,  352;  titnoriensis , 

46;  xanthopygus,  351 
Cestracion,  63 

Cetaceans  of  American  Tertiary,  68 
Cetotherium,  68 
Ceylon,  when  separated  from  India, 

293 

Ceylonese  sub-region,  266 
Chcerokyus,  376 
Chceromeryx,  203 
Chtzropotamida:,  161,  192 
Cheer opotamus,  161,  192 
Cheer optis,  39 

Characiniidce,  distribution  of,  134 
Chcetomys,  go,  136 
Chalicomys,  159,  194,  196,  313 
Chalicotheriidce,  196,  200 
Chalicotherium,  170,  192,  200 
Chameleons,  distribution  of,  222 
Chamois,  324 
Chelonians  of  Notogsea,  62 
Chevrotains,  164,  281 
Chimarrogale,  272 
Chilian  sub-region,  135 
Chimpanzees,   180,  231,  256 
Chinchillas,  89,  136 
Chinchillidee,  89 
Chipmunks,  313 
Chiromyidce,  181 
Chiromys,  219 
Chiropodomys,  278 


INDEX. 


389 


Chiroiiectes,  107 

Chiru,  325 

Chirnromys,  41 

Chlamydophorus,  94,  96,  137 

CJilamydotherium,  96 

CholcepuS)   10 1 

Chriacidie,  371 

C/iriacis,  373 

Christmas  Island,  303 

Chromididiz,  134 

Chrotomys,  277 

Chrysochloridiz,  230,  234 

Ckrysochloris,  230,  234 

Civets,  182;  Oriental,  273 

Clccnodon,  159 

Claviglis,  238 

Coatis,  72,  365 

Cobus,  200,  206,  -245 

Cwlogenys,  89,  136 

Colics,  254 

Collide?,  254 

Colobus,  231 

Colymbus,  347 

Comoro  Islands,  213 

Conepatus,  73,  365 

Condylura,  341 

Conihtrus,  41 

Connection    between   Africa   and    S. 

America,    127 ;    Australia   and   S. 

America,  125 
Connochcetes,  244 
Continental  islands,  10 
Continents,  permanency  of,  22 
Cooke,  Mr,  on  S.  American  molluscs, 

124;  on  Antillean  molluscs,    139; 

on   N.    American   land    Mollusca, 

3n 
Cope,    Prof,  on  N.   American   fossil 

cetaceans,  68 
Coral  his,  132 
Coryphodontidce,  174 
Coryphodon,  174,  378 
C0.SWJ.*-,  368 
Cotton-rat,  88 
Cotylopidce,  373 
Cotylops,  375 
Coypu,  91,  137 
Crateromys,  277 
Cricetince,  160,  183,  239 
Cricetomysy  239 
Cricetus,  160,  192,  196,  322 
Crocidura,  195,  272 
Crocodilus  porosus,  62 
Crossarchus,  235 
Crossopus,  319 
Cryptobranchus,  35 
Cryptodira,  distribution  of,  8 


Cryptoprocta,  220 
Ctenodactylidcz,  90 
Ctenodactylus,  91,  212,  240,  323,  357, 

358 

Ctenomys,  90 
Cuniculus,  314 
Cuscuses,  36,  46 
Cusimanse,  235 
Cycloturus,  102 
Cyncelurus,  202,  234,  249,  272 
Cynic tis,  235 
Cynodictis,  158,  191 
Cynodon,  191,  193 
Cynopithecus  niger,  47 
C>»,  235,  274 
Cynogale,  273 
Cynomys,  342 

Dacrytherium,  185,  192 

Dactylomys,  91 

Dactylopsila,  37 

Dcedicurus,  98 

Damaliscus,  244 

Damuda-Talchir  flora,  153 

Dapedoglossus,  134 

Daphcenus,  158 

Dasornis,  60 

Dasymys,  238 

Dasyprocta,  98,  137 

Dasyproctida,  89 

Dasypodidce,  94 

Dasypus,  94 

Dasyuridce,  39  ;  Neogaeic,  108 

Dawkins,   Prof.  B.,  on  the  range  of 

the  mammoth,  334 
Decastis,  no 
Deer,   164,  267,  367  ;  Oriental,  280 ; 

their     absence     from      Ethiopian 

Africa,   230,  258 
Deep-sea  deposits  in  islands,  23 
Degu,  90 
Dendrogsea,  26 
Dendrogale,  270 
Dendrolagus,  36 
Dendromys,  238 
Deomys,  239 
Desman,  321 
Deserts  as  barriers,  15 
Diadiaphoriis ,  80 
Diatryma,  60 
Dichobumts,  163,  192 
Dichodon,  192 
Dichodontidce,  185,  192 
Dicroceros,  196 
Dicotyles,  73,  368 
Dicotylida,  74,  375 
Dicynodonts,  151 


390 


INDEX. 


Didelphyidce,  Neogseic,  107 
Didelphys,    51,    roy,    192,    193,    194, 

368 

Dingo,  42 
Dinocyon,  158,  195 
Dinomyidce,  89 
DinornithidfB,  58 
Dinosauria,     151  ;    wide    distribution 

of,  8 

DinotheriidcB)  187 

Dinotherium,  173,  187,  196,  200,206 
Dist<echurus,  37 
Dispersal,  barriers  to,  10 
Distribution  of  groups,  9 
Distributional  Areas,  19 
Distribution  of  families,  20  ;  genera, 

19  ;  orders,  21  ;  species,  19 
Diplomesodont  320 
Dipodidce,   240,    342  ;    not  Hystrico- 

morpha,  159  ;  E.  Holarctic,  322 
Dipodomys,  367 
Diprotodon,  38 
Diprotodonts,  34 
Dipus,  322 
Diver,  347 

Dobson,  Dr,  on  Dipodidiz,  159 
Dolichotis,  88,  136 
Dolichopithecus,  180 
Dorcatherium,    164,    186,    196,    197, 

198,  204,  242,  256 
Dorcopsis,  36 
Dremotherium,  194 
Dromatherium,  54 
Dormice,  322  ;  African,  238 
Dorcatragus,  245 
Dromicid)  37 
Dromiciops,  107 
Dryolestes,  54 
Dryopithecus,  180 
Dual  origin  of  groups,  380 
Duckbill,  32 
Duikerboks,  244 

Earth-worms,  126;  of  Patagonia  al- 
lied to  those  of  Australasia,  126 

East  Central  African  sub  -  region, 
261 

Eastern  Arctogsea,  179;  its  Tertiary 
mammalian  fauna,  190 

Echidna,  33 

EchidnidcBi  33 

Echinomys,  91,  136 

Echinops,  220 

Echinothrix,  48 

Edentata,  92 

Edentates,  doubtful  fossil  forms,  187 

Effodientia,  187,  192 


Elands,  247 
ElaphodiiS)  326,  356 
Elasnwtherium,  333 
Elephas,  172,  206,  247,  282 
Elephants,     172  ;      Stegodont,     172  ; 

African,  247  ;  dwarf  Maltese,  339  ; 

European  Plistocene,  334  ;  Japan- 

ese fossil,  356 
Elephantida:,  Oriental,  282 
Elk,  164,  315 
El  lolnus,  322 

Elotherium,  161,  192,  193 
Emballonuridce,  S.  American,  70 
Enhydriodon,  195,  203 
Endrina,  217 
Eocardia,  91 
Eocardiidce,  91 
Eodidelphys,  107 
Epanorthidce,  no 
Epanorthus,  no 
Epomopfioriis,  221,  232 
Eqttida,  165  ;  S.  American,  75  ;  Ori- 

ental, 282 
Equus,  1  68,  206,  247,  282,  359;  its 

dual  origin,  380 
Erethizon,  90,  343 
Ericulus,  220 
Erinaceida,  181,  271 
Erinaceus,  271,  233 
Eriodes,  136 
Eriomys,  89,  136 
Ethiopia  and  India,  their  connection, 

257 

Ethiopian  Mammals,  230 
Ethiopian  Region,  25,  227  ;  its  past 

history,  255 
Eucardiodon,  89 
Eucholczops,  107 
Euchoretes,  322,  351 
Eumeces,  357 
Eupetaurus,  322 
En  pier  es,  220 
European  sub-region,  348 
Etismilus,  157,  191 
Eiitat2is,  94 
Eutheria,  34 

Everett,  Mr,  on  Malayan  fauna,  294 
Extinct  mammals  of  Western  Arcto- 

g«a,  371 
Extinction  of  large  mammals,  18 


347 

Falkland  Islands,  140 
Fallow  deer,  326,  359 
Faunas,  community  of  early,  148 
Felidcc,  African,  234,  249  ;  Holarctic, 
312  ;  Neogaeic,  71  ;  Oriental,  272 


INDEX. 


391 


Felis,    198,    202,    249;   concolor,   71; 

MO**/,  351  ;  megalotiS)  46 
Fernando  Noronha,  140 
/W«?r,  342 
Fish-eating  rats,  88 
Fishes,  community  between  those  of 

Australasia  and  S.  America,  125 
Flora,  community  of  Southern  Palseo- 

zoic,  153 

Flying  phalangers,  37 
Flying-squirrels,  African,  237 
Forbes,  Dr,  on  Antarctica,  128,  129 
Forest-bed,  its  fauna,  335 
Fossa,  220 
Foxes,  317 

Galago,  217,  231 

Galaginft,  217 

Galapagos  Islands,  140  ;  Tortoises  of, 

141 

GaleopitheciiS)  268,  270 
Galerisctts,  237 
Galerix,  197,  270 
Galictis,  73 
Galidia,  220 
Galidictis,  220 
Garzoniidce,  1 1  r 
Gastornis,  60 
Gaur,  278 
Gazella,    199,    206,    245,    325,    358; 

gutturosa,  351 
Gazelles,  245 
Ge/oats,  192,  193 
Gemsbok,  245 
Genetta,  182,  321,  358 
Genets,  182 
Geogale,  219 
Geographical  changes  in  Europe  since 

the  Plistocene,  337 
Geological  sequence,  7 
Gcomytd&i  366 
Geomys,  366 
Georhychus,  239 
Gerbillince,  183 
Gerbillus,  357 
Giant  Salamander,  356 
Giant   birds   of   Patagonia,   60;    the 

Eocene,  60 
Gibbons,  180,  269 
Giraffa,  186,  189,  204,  242 
Giraffidce,  186,  242 
Glacial  epoch,  9;  period,  in  America, 

345 

Glossotherium,  105 
Glyptodon,  96 
Glyptodontidce,  96 
Gnus,  244 


Goats,  324 

Golunda,  278 

Gorilla,  180,  231 

Graphiurus,  238 

Greenland  Falcon,  347 

Gregory,  Dr,  on  communication  be- 
tween Atlantic  and  Pacific,  1 39 ;  on 
modern  origin  of  Atlantic,  23,  135 

Grison,  73 

Groove-toothed  mice,  88 

Ground-sloths,  102 

Guanaco,  74,  136 

Gundi,  357 

Gymnura,  271 

Gymnobelideus,  37 

Guto,  312 

Habrocoma,  90 

Hamsters,  160,  322 

Hapalemur,  219 

Hapalidtz,  69 

Hafalomys,  278 

Hapalotts,  41 

Haploceros,  165,  343 

Haplodon,  342 

Harnessed  Antelopes,  246 

Harpyia,  42 

Hawaiian  region,  27,  30,  45 

Hedgehogs,  181 

Hedley,  Mr,  on  the  isolation  and  con- 
nections of  Australia,  59 

Hegetotherium,  85 

Heilprin,  Dr,  on  migrations  of  mam- 
mals, 15;  on  zoological  regions,  26, 
29 ;  on  the  Boreal  Region,  346 ; 
on  the  Boreal  Fauna,  347 

Helaletes,  165 

Helicophora,  200 

Helictis,  276,  355 

Helladotherium,  186,  198 

Helmsley,  Mr,  on  Galapagos  flora, 
142 

Helogale,  235 

Hemicentetes,  220 

Hemicyon,  195 

Hemigale,  273 

Hemigalidia,  220 

Hemimeryx,  203 

Hemipsalodon,  258 

Hemitragus,  206,  209,  230,  279,  288, 
324 

Herpestes,  182,  235,  321,  358,  360 

Heteromys,  136,  367 

Himalayan  sub-region,  266 

Hipparion,  167,  200,  206,  247 

Hippidiuni,  75 

Hippohyiis,  203 


392 


INDEX. 


Hippopotamida,  184,  241 

Hippopotamus,  204,  241,  256;  cross- 
ing from  Africa  to  Madagascar,  223 

Hippoligris,  247 

Hippotragus,  199,  206,  245 

Holarctic  region,  26,  27;  physical 
features  and  extent,  309 ;  character- 
istics of  the  fauna,  311;  its  unity, 
316;  eastern  division,  319;  Plisto- 
cene  Fauna  of  the  eastern  division, 
328 ;  western  division,  340 

Holochilus,  88 

Homacodon,  163 

Homalodontotheriunt)  82,  170 

Homunculus,  69 

Hornbills,  254 

Horse,  317 

Horses,  S.  American,  75 

Hose,  Mr.,  on  Malayan  fauna,  294 

Howorth,  Sir  H.  H.,  on  the  mixture 
of  northern  and  southern  types  of 
mammals  in  the  Plistocene,  329 

Humidity,  influence  of,  5 

Hundes,  mammalian  remains  in,  354 

Hunting-dog,  236 

Hunting-leopard,  249,  272 

Hildas,  91,  137 

Huxley,  Prof.,  on  zoological  regions, 
25,28;  on  northern  origin  of  south- 
ern birds,  130 

Hyana,  182,  198,  235,  249,  273,  321 

HycBnarctus,  158,  195,  198,  202,  321 

HyaenictiS)  182,  198 

Hycenidce,  an  Old  World  group,  182 

Hycenodon,  158,  192,  193,  203 

Hy&nodontida,  373 

Hydaspitherium,  186,  204 

Hydraspis,  132 

Hydrochcerus,  88,  136 

Hydromyin<z,  40 

Hydromys,  40 

Hydropotes,  326,  356 

Hylobates,  180,  269 

Hylomys,  271 

Hyopotamus,  161 

ffyopsodtts,  155 

Hyotherium,  194,  196,  203,  376 

Hyperodapedon,  63 

Hypertragulus,  164 

Hypocetus,  68 

Hypselornis,  60 

Hypsipetes,  254 

Hyraces,  248,  268 

Hyrachyus,  166,  192 

Hyracodon,  no,  170 

Hyracoidea,  relationship  to  toxodonts, 
85,  248 


Hyracotherium,  161,  165 
Hystricomorpha,  their  abundance  in 

Neogaea,  88 
Hystricidce,    S.  American,    90;    Ori- 

ental, 278 
Hystricinte,  184 
Hystrix,  196,  198,  203  ;  javanica,  46 

Ibex,  359 
Iceland,  350 
IchthyomyS)  88 

Icticyon,  72,  136 

Ictitherium,  182,  198 

Ictonyx,  236 

Idiurus,  238 

Iguanidce,    distribution,     132;    fossil, 

133 
Iguanadon,  151 

Iguanas,  fossil  in  Europe,  62;  of  Fiji 

and  Friendly  Islands,  62 
India  and  Ethiopia,  their  connection, 

209,  257 

Indian  region,  25  ;  sub-region,  266 
Indo-Malayan  sub-region,  266 
Indo-  African  region,  26 
Indo-Chinese  sub-region,  266,  267 
Indrisince,  217 
Indrodon,  373 
Introduced  mammals,  17 
Insectivora,  Neoggeic,  70;  Arctogaeic, 

156;  Holarctic,  312;  North  Amer- 

ican, 341  ;  Sonoran,  364 
Ireland,  its  connection  with  Britain, 

340 

Irish  deer,  326,  333 
Irrisoridce,  254 
IsectolophuS)  165 

Islands,  continental,  60;  oceanic,  10 
Isomys,  239 
IssiodoromyS)  89,  92 
Ixocincla,  254 

Java,  distinctness  of  its  fauna,  300 
Jerboa-rats,  41 
Jumping-mice,  342 
Jungle-cat,  249 

Kangaroo-rats,  366 

Kangaroos,  35 

Kinkajou,  72 

Kirby,  Mr,  on  Holarctic  Lepidoptera, 

310 

Kudus,  246 
Kurtz,  Dr,  on  Argentine  fossil  flora, 


Labyrinthodontia,  152 


INDEX. 


395 


Lagidium,  89,  136 

Lagomyidce,  159,  314 

Lagomys,  196,  314 

Lagopus,  347 

Lagorchestes,  36 

Lagostomatidce,  89 

Lagostomus,  89,  136 

Lagos  tr op  hits,  36 

Lagothrix,  136 

Lama,  74,  136,  377 

Lamprotornis,  254 

Langiirs,  269,  360 

Laniarius,  254 

Lanthanotheriiun,  195,  197,  270 

Zate*,  312,  356,  347 

Le  Conte,  Dr,  on  separation  of  N.  and 
S.  America,  118 

Leberon  Fauna,  197 

Leith-Adams,  Dr,  on  the  Maltese 
Islands,  337 

Lcithia,  333 

Leopard,  249 

Lepidosiren,  133 

Lepidosirenida,  distribution  of,  133 

Leporida,  160;  Holarctic,  314;  Neo- 
gaeic,  92 

Leptarctiis,  72 

Leptodon,  200 

Leptomanis,  187,  192 

Leptomeryx,  164 

Leptotragtdus,  163,  377 

Lepus,  203 

Lemmings,  314 

Lemuroids,  181,  191;  Arctogaeic,  155; 
character  of,  216;  early  entrance 
into  Ethiopian  and  Malagasy  re- 
gions, 223;  extinct,  371 

Leimiridce,  181,  217,  270 

Lemur,  217 

Libytherium,  186 

Limacomys,  238 

Linsanga,  235,  256,  273 

Lion,  249,  272 

Listriodon,  196,  203 

L^thocran^^ls,  245 

Litopterna,  75 

Lomaphorus,  28 

Lomvia,  347 

Loncheres,  91,  136 

Lophiodontidce,  165,  192 

I^ophiodon,  165,  187 

Lophiomys,  239 

Lophiomeryx,  192 

Lophuromys,  239 

Z0rw,  256,  270 

Loxomylus,  87,  137 

Lutra,  202 


Lycaon,  236 
Lynxes,  312,  317 

Macacus,    180,    202,    269,    355,   357,. 

360  ;  cynomolgus,  46  ;  maurus,  47 
Machcerodus,  157,195,202;  Neogseicr 

71 

Macropus,  35 

Macrauchenia,  77 

Macropodidce ,  35 

Macroscelides,  233,  357 

Macroscelididce,  233,  268;  fossil,  191 

Macrotherium,  172,  196,  200 

Madagascar,  213 

Madoqtia,  244 

Malabar  sub-region,  266 

Malacomys,  238 

Malagasy  region,  26,  213;  mammals, 
214;  molluscs,  223 

Malayan  sub-region,  267,  294 

Malaysia  and  W.  Africa,  affinity  of 
faunas,  292 

Malta,  part  of  the  mainland,  337 

Mammals,  the  oldest,  9;  importance 
of,  9 ;  classification  of,  9 ;  swimming 
powers  of,  13;  means  of  dispersal  of, 
13;  introduced  by  man,  17;  extinc- 
tion of,  18;  of  Oriental  region,  267 

Mammoth,  317 

Man-like  apes,  distribution  of,  180 

Manidce,  187;  Oriental,  283 

Mam's,  187,  249,  284 

Man's  influence  on  distribution,  16 

Maraga  Fauna,  197 

Marmots,  267 

Marmosets,  69 

Marsupial  mole,  40 

Marsupialia,  33 

Marsupials,  33;  Notogseic,  34;  pa- 
Iseontological  history  of,  50  ;  fossil, 
51;  survival  in  Asia,  55,  57;  date 
of  migration  into  Notogaea,  60; 
Neogaeic,  107 

Mantchurian  sub-region,  355 

Martens,  318 

Mastacomys,  41 

Mastodon,  196,  200,  206,  172;  S. 
American,  86 

Mauritius,  213 

Medio-Columbian  region,  26 

Mediterranean  region,  26,  319;  sub- 
region,  357 

Megaladapis,  218 

Megalobatrachus,  356 

Megalonyx,  105 

Megalotheriida,  102 

Megalotherium,  103 


394 


INDEX. 


Metes,  321 

Mellivora,  202,  237,  256,  276 

Mehirsus,  202,  274 

Mentawi,  303 

Mephitis,  73,  365 

Mergulns,  347 

Merriam,  Dr,  on  effects  of  tempera- 
ture, 4;  on  W.  Indies,  140;  on  a 
Boreal  region,  309;  on  the  effects 
of  a  glacial  period  in  N.  America, 
345 ;  on  the  Transition  zone,  360, 
361 ;  on  the  Sonoran  region,  363 

MerioneS)  357 

MerycopotamuS)  203 

Mesohippus,  167 

Mesomys,  91 

Mesonychidce,  371 

Mesonyx,  373 

MesoptthecW)  1 80,  197,  200 

Mesosaurus^  133 

Metatheria,  33 

Mexican  sub-region,  136 

Miacidce,  373 

MicroMotheriida,  109 

Microbiotheriu  m,  109 

Microgale,  220,  234 

Microchcerus,  155,  156,  181,  191 

Micropholis,  152 

Micropus,  352 

Microtinat)  160 

Microtus,  267,  313,  318 

Minas  Geraes,  its  caves,  66 

Mixodectes,  373 

Mixodectidtz,  373 

Moas,  58 

Moles,  321 

Mole-rat,  322,  358 

Mole-voles,  322 

Moluccas,  46 ;  their  isolation,  49 

Mongolian  Pliocene,  201 

Monkeys,  S.  American,  69 ;  distri- 
bution and  dual  origin  of,  179,  180 

Monte  Hermoso,  its  mammaliferous 
beds,  67 

Monotremata,  32 

Monotremes,  32  ;  date  of  migration 
into  Notogaea,  60 

Morenia,  91 

Morosaurus,  152 

Moschus,  204,  325 

Mouflon,  358 

Mountains  as  barriers,  15 

Mule-deer,  368 

Multituberculata,  33 

Muntjacs,  281 

Muridce,  160,  183;  Notogaeic,  41; 
African,  238;  Holarctic,  313;  E. 


Holarctic,  322  ;  N.  American,  342  ; 
Sonoran,  365 
Murince,  160;  an  Old  World  group, 

183 
Muscardinus,  322 

Musk-deer,  325 
Musk-kangaroo,  36 
Musk-ox,  315,  318,  344 
Musk-rat,  342,  366 
Musk-shrews,  272 
Musophagidce,  254 
Mustela,  195,  198,  202,  236,  358 
Mustelzdce,  236;  Neogseic,  73;  Sono- 
ran, 365 

Mus,  Notogaeic  species,  41 
Mus  armandvillei,  46 
Mydaus,  276 
Mylodon,  103 
My  odes,  314 
Myogale,  321 
Myolagus,  196 
Myopotamus,  91,  136 
Myoscalops,  239 
Myosorex,  233 

Myoxidce,  322  ;  African,  238 
Myoxus,  192,  196,  322 
Myrmecobius,  40 
Myrmecophaga,  102,  136 
Mynnecophagidce,  101 
Mystromys,  238 
Myxopoda,  132 

Nandinia,  235,  256 

Nannosciurus,  238,  277 

Nanotragus,  244 

Nasalis,  270 

Nasua,  72,  365 

Nearctic  region,  25 

Necrogymmirus,  182,  191,  271 

Nectarinia,  254 

Necrolestes,  71 

NecrotnaniS)  187,  192 

Nemorhtedus,  280,  324 

Neofiber,  366 

Neogaea,  25,  27,  64;  its  mammali- 
ferous deposits,  66  ;  its  early  fauna 
distinct,  \\.i\  distinctness  of  its 
modern  fauna,  123;  its  peculiar 
birds,  123;  its  land-molluscs,  124; 
connected  with  Notogaea,  125;  con- 
nected with  Afi-ica,  127 

Neogseic  fauna,  origin  of,  124 

Neogaeic  realm,  27, 64 ;  its  sub-regions, 

»35 

Neoplagiaulax,  156 
Neoremys,  91 
Neotoma,  342,  366 


INDEX. 


395 


Neotragus,  244 

Neotropical  region,  25,  27,  64 

Nesocerodon,  92,  89,  192 

Nesocia,  203 

Nesodon,  83 

Nesotragus,  245 

Neumayr,  Dr,  on  a  land-bridge  across 
Atlantic,  133 

NeurotricJnis,  312 

New  Guinea,  its  mammals,  31 

New  Siberian  Islands,  348 

New  Zealand  mammals,  45 

New  Zealand,  its  relations  with  Aus- 
tralia, 58,  59 

Nimravus,  157 

North  America,  affinity  of  its  fauna 
with  that  of  E.  Asia,  57 ;  its  relation 
to  Asia,  310 

North  American  fauna,  340;  Tertia- 
ries,  sequence  of,  375 

Nothropus,  107 

Nothrotherium,  107 

Notiosorex,  364 

Notogcea,  25,  27,  28;  its  isolation 
comparatively  modern,  57,  58 ;  its 
chelonians,  62 ;  its  right  to  form  a 
realm,  62 ;  northern  origin  of  its 
fauna,  62 

Notogseic  Realm,  27,  28 

Notary ctes,  40 

Notoryctida,  40 

Nototheriuni)  38 

Novo-Zelanian  region,  25 

Nyctea,  347 

Nyctereutes,  355 

Nycticebus,  256,  270 

Oceanic  islands,  10 

Oceans,  permanency  of,  22 

Octodon,  90 

Octodontidce,  137,323;  Neogaeic,  90; 

African,  240 
Oligocene  fauna,  190 
Onohippidium*  75 
Onychogale,  36 
Onychomys,  366 
Opossums,  368;  European  fossil,  51  ; 

Neogaeic,  107 ;  originally  a  northern 

group,  1 08 
Orangs,  180,  268 
Oreodonts,  373 
Oreopithecus,  181 
Oreotragus,  245 
Or  ibid,  245 
Orias,  199,  206,  247 
Oriental  Region,  25,  27,  264;  its  past 

history,  288 


Ornithorhynchus,  32 

Ornithorhynchida,  32 

Ornithogaea,  26 

Orthomys,  91 

Orycteropodidce,  187,  249 

Orycteropus,  187,  200,  249,  268 

Oryx,  199,  245,  358 

Oryzomys,  366 

Oryzorictes,  220 

Osborn  and  Earle,  Messrs,  on  Puerco 

Fauna,  154 

OsteoglossidcE,  distribution  of,  134 
Osteoglossum,  134 

Ostriches,  255,256;  distribution  of,  129 
Otocyon,  202,  236 
Otomys,  238 
Ovibos,  165,  315,  344 
Ovt's,  165,  280,  314,  324,  358 
Ovis  antmon,    352 ;    hodgsoni,    352  ; 

poll)  352 

Oxycena,  192,  373 
Oxyodontotherium,  79 

Pacas,  89,  136 
Pachynolophus ,  166,  192 
Pachyruchida,  85 
Pachyruchus,  85 
Pachyuromys,  238,  239 
Pala,  245 

Palsearctic  region,  25,  26 
Palcecerinaceus,  182 
Palaeogsea,  25 
Palceomanis ,  187 
PalcEomephitis,  198 
Palceomeryx,  196,  204 
Palaeontology,  importance  of,  6 
Palaoprionodori)  191 
Palceorias,  199 
Palaorycteropus,  187,  192 
Palczosyops,  170,  377 
Palawan  group,  294 
PalcEotapirus,  165 
Palaotheriidce,  166,  192 
Palaotherium,  166,  187,  192,  193 
Palaotragus,  186,  199 
Palhyana,  182,  198 
Palorchestes,  36 
Pampas,  66 
Pampean  beds,  66 
Panda,  212,  275 
Pangolins,  187,  249,  283 
Panochthus,  98 
Pantholops,  324 
Papio,  1 80,  202,  231,  256 
Paracetus,  68 

Paradoxurus,   256,  273  ;  hermaphro- 
ditus,  46  ;  muschenbroecki,  47 


396 


INDEX. 


Parana  beds,  88 

Patagonian  beds,  67  ;  cavy,  136 

Patriofelis,  373 

Peccaries,  74/368,  375 

Pectinator ,  91,  240 

Pedetes*  240 

Pelea,  245 

Pellegrinia,  91,  212,  240,  323 

Pelomedusa,  131 

Pelomedusidce,    distribution    of,    131, 

132 

Peltephilus,  96,  112 
Pelycodus,  155 

Penguins,  a  southern  group,  131 
Peragale,  38 
Peramelidce,  38 
Perameles,  38 
Peratherium,  51 
Perimys,  90 

Perissodactyles,  Arctogseic,  165 
Permanency  of  continents  and  oceans, 

22 

Perodicticus,  231,  256,  270 

Perodipus,  367 

Perognathjts,  367 

Petauroides ,  37 

Petaurus,  37,  46 

Petrogale,  36 

Peiromys,  240 

Phacochcems,  185,  203,  242,  268 

Phalanger^  36,  46 

Phalangeridie,  36 

Phalangers,  36 

Phascologale,  39 

Phascolomyidcc,  38 

Phascolomys,  38 

Phascolonus,  38 

Phascolotherium,  52 

Phenacodus,  166,  174 

Phenacomys,  342 

Philippine  sub-region,  267 ;  its  fauna, 

304 

Phlczomys,  277 
Phororhachis,  60 
Phosphorites,  191 
Phyllostomatidce,  70 
Picas,  314 

Pichiciagos,  94,  96,  137 
Pikermi  Fauna,  197 
PithechiruS)  278 
Pithed  a,  136 
Plagiaulax,  147 
Plagiodon,  91,  137 
Plantain-eaters,  254 
Platanistid<z,  112 
Plattmys,  132 
Platycercomys,  322 


Pkctrophanes,  347 
PlesiarctotnyS)  160 

Plesictis,  191,  193 

Plesiospermophilus,  192 

Plesiorycteropus,  249 

Pleurodira,  distribution  of,  8  ;   131 

Plexoch&rus ,  89 

Pliauchenia,  163,  377 

Pliocene  Fauna  of  Arctogsea,  197 

Pliolagostomus,  89 

Pliopithecus,  180 

Plohophorus,  98 

Podocnemis,  131 

Poebrotheriuni)  1 63 

Pcecilogale,  236 

Poiana,  235,  256 

Poly  mastodon,  149 

Polynesian  region,  27,  29,  45 

Polyprotodonts,  34 

Pontoporia,  112 

Pouched  mice,  39 

Pouched  rats,  136,  366 

Porcupines,  184  ;  S.  American,  90 

Potamogale,  234 

Potamogalidcc,  220,  234 

Pot  or ous,  36 

Pottos,  231,  256,  270 

Prehensile-tailed    mammals   in    Neo- 

gsea,  124 
Priodon,  94,  136 
Proboscideans,  Neogaeic,   86  ;  Arcto- 

goeic,  172 

Proboscis  Monkey,  270 
Procamehts,  163,  377 
Procavia,  248 
Procaviidce,  248,  268 
Procoptodon,  36 
Procyon,  72 
Procyonid(Z,     364  ;      Neogseic,      72 ; 

Oriental,  275 
ProdidelphyS)  107 
Prodremotherium,  164,  192 
Proechidna,  33 
Progenetta,  195 
Prolagostomus,  89 
Protneles,  198 
Prongbuck,  367 
PropalceohoplophontS)  i  oo 
Protalpa,  156,  321 
Protapirus,  165,  192 
Protechinomys,  1 92 
Proteleidce,  235 
Proteles,  235 
Proterotheriidce,  80 
Proterotherium,  80 
Prothylacinus,  40,  108 
Protoceras,  376 


INDEX. 


397 


Protoceratida,  376 
Protogonia,  174 
Protohippus,  168 
Protolabis,  163 
Protopterus,  133 
Ptotoreodon,  375 
Prototheria,  32 
Protragoceros,  199 
Proviverra,  159,  192,  193 
Pseudalurus,  195,  198 
Pseud h  apalops,  \  o  7 
Pscitdochirus,  37 
Psendocyon,  195 
Pseudorhynchocyon,  191,  233 
Pseudoscittrus ,  192 
Psittaci,  distribution  of,  129 
Psittacotheriidce,  379 
Psiltacotherium,  379 
Psittacula,  distribution  of,  130 
Ptarmigan,  347 
Pterodon,  158,  192,  193 
Pteromys,  268,  277 
Pteropodidce,  Notogseic,  42  ;  distribu- 
tion of,  221,  225,  232 
Pteropiis,  221,  232,  268 
Ptilocercus,  270 
Puerco  Fauna,  153 
Puma,  71 
Pyrotheria,  85 
Pyrotherium,  85 

Quagga,  247 

Quercy  Phosphorites,  191 

Raccoons,  72,  364 

Rafts,  15 

Rangifer,  164,  315 

Raphiceros,  245 

Rat-kangaroos,  36 

Ratitse,  distribution  of,  129 

Ratite  birds,  their  migration,  58,  60 

Realms,  25,  27 

Regions,  25,  27 

Reindeer,  164,  315 

Reptiles,  antiquity  of,  8 

Ratels,  237,  256 

Rhea,  130 

Rheebok,  245 

Rhinoceroses,  169;  African,  247,  256; 

European  Plistocene,  333 
Rhinoceros,  169,  192,   193,  206,  247, 

256,  281 

Rhinocerotutei  169;  Oriental,  281 
Rhithrodon,  88,  140,  365 
Rhithrodontomys,  365 
Rhithrosciurus,  277 
Rhizomys,  183,  203,  240 


Rhynchocyon,  233 

Rhynchomys,  278 

Rhynchosaurns,  63 

Rietbok,  245 

Rio  de  la  Plata,  its  characters,  67 

River-channels,    buried,  W.    Indian, 

.'38 

Rivers  as  barriers,  14 
Rocky  Mountain  Goat,  343 
Rodents,  Arctogseic,  159;  Ethiopian, 

237  ;  Holarctic,  313;  Neogseic,  86  ; 

North  American,   342 ;    Notogaeic, 

41  ;  Oriental,  277  ;  Sonoran,  365 
Riipicapra,  324 
Range  of  Carnivora,  5 
Ruscinomys,  91 

S.  American  Tertiaries,  their  age,  68 
S.  America  connected  with  Australia, 

125;  connected  with  Africa,  127 
S.   American   element   in    Ethiopian 

fauna,  128 
Sable  Antelope,  245 
Sab  re- tooths,  Neogaeic,  71 
Sahara,  its  action  as  a  barrier,  259 
Saharan  sub-region,  260 
Saiga,  324 
Sam  bar,  280 
Samos  Fauna,  197 
Samotheriiim,  186,  198 
Sandwich  Islands,  45 
Santa    Cruz    beds,    67 ;     fauna,    its 

origin,   124 
Sarcophilus,  39 
Seal  ops,  364 
Scapanus,  364 
Scaptochirus,  321 
Scaptonyx,  321 
Scelidotheriuin,  105 
Scharff,  Dr,  on  the  origin  of  the  Irish 

fauna,  340 
Schizodon,  91 
SciuridcE,  Holarctic,  313 
Sciuroides,  192 
SciurofteruS)  195,  277 
Sciurus,  195,  277  ;  prevosti,  48 
Sclater,  Dr,  on  zoological  regions,  25, 

26;  on  the  antelopes  of  Somaliland, 

261 

Scleromys,  91 
Scotonycteris,  221,  232 
Scott,  Prof,  on  origin  of  Canis,  158  ; 

on  Dipodidce,  159;  on  the  Equidtf, 

168 

Sea-otter,  312  ;  356 
Secondary  Reptiles,  wide  distribution 

of,  151 


398 


INDEX. 


Secretary-birds,  254 

Selvas,  in 

Semioptera,  48 

SemnopithecuS)   180,   202,    269,    357, 

360 

SerpentariuS)  254,  255 
Sewellels,  342 
Seychelles,    213;    indicate   a  line  of 

connection     between     India     and 

Africa,  224 
Sheep,  324 
Shrews,  267,  272 
Side-necked  tortoises,  distribution  of, 

131*  13* 

Siderolites,  191 

Sifaka,  217 

Sigmodotii  88,  366 

Szmia,  180,  202,  268 

Simiidcz,  268 

Simocyon,  198 

Siphneus,  322 

Sitomys,  88,  137,  160,  322,  342,  365 

Siwalik  Fauna,  201 

Sivatherium,  186,  204 

Skunks,  73,  365 

Sminthopsis,  39 

Sminthus,  322,  359 

Snow-bunting,  347 

Snowy  Owl,  347 

Solenodon,  70,  138 

Solenodontida,  70 

vSomaliland,  its  fauna,  261 

Sonoran  region,  26,  27,  363  ;  its  dis- 
tinctness, 380 

Sorex,  191,  198,  267,  272,  312 

Soricidce,  136,  156,  319;  Ethiopian, 
23°»  233  >  Holarctic,  312;  So- 
noran, 364 

Soriculus,  272 

South  American  region,  26 

South  America,  separated  from  N. 
America,  117;  invaded  by  northern 
animals  in  the  Pliocene,  119 

Southern  Pateozoic  Continent,  153 

South  African  sub-region,  261 

Spalacidce,  183,  239,  322 

Spalacopus,  91 

Spalacotheriidce,  $2 

Spalax,  183,  322,  358 

Spencer,  Mr,  on  buried  West  Indian 
river-channels,  138 

Spermophilus,  159,  351 

Spheniscida,  a  southern  group,  131 

Sphenodon,  8,  63 

Sphodromys,  90 

Spiny  anteater,  33 

Spiny  rats,  136 


Spilogale,  365 

Sparassodonta,  108 

Squirrels,  Flying,  237 

Star-nosed  mole,  341 

Station,  2 

SteatomyS)  239 

Stegodont  elephants,  206 

Stenodelphis,  1 1 2 

Stenoplesictis ,  182,  191 

Stereornithes,  60 

Stereo  stern  um ,  133 

Sternothoerus*  131 

Sthenurus,  36 

StiromyS)  90 

Stonestield  slate,  its  marsupials,  52 

Strepsiceros,  199,  206,  246 

Strut hio,  255;  distribution  of,  129 

Submerged  forests  of  Britain,  339 

Sucker-footed    bats,    distribution    of. 

13 r 

Suida:,  184,  375;  Oriental,  281 
Sumatra,  300 
Siiricata,  235 
Sus,  184,  203,  242,  267,281;  absence 

of,  in  Ethiopian  region,   230  ;  cele- 

bensis,  48 ;  papuensis,  42 
Swamp-deer,  280 
Swayne,   Capt.,   on  the  fauna  of  So- 

maliland,  261 
Syria,  formerly  connected  with  Africa, 

259 
Systemodott)  165 

Takin,  324 

Talpa,  156,  191,  271,  321 

Talpidce,   156,  271,   312;    absence  in 

Ethiopia,  230;  Sonoran,  364 
Tamandua,  102 
Tamarao,  47,  279 
Tamias,  159,  313 
Tatniascvurus )  344 
7\ipiridce,  165;  Oriental,  282 
Tapirs,  74 

Tapirus,  73,  165,  194,  282 
Tarsiidce,  181,  270 
Tarsipes,  37 

TarsiuSi  2  70 ;  fuscomanus,  47 
Tasmania,  31 
Tasmanian  devil,  39 
Tatusia,  94,  368 
Tayra,  73 
Tchitrea,  254 
Telephomis,  254 
Temnocyon,  158 
Temperature,  influence  of,  3 
Tenrec,  220 
Teonoma,  344,  366 


INDEX. 


399 


Tertiary  mammalian  fauna  of  Eastern 

Arctogoea,  190 
Tesiudo,  distribution  of,  129 
Tetraceros,  206,  244,  280 
Tetraconodon,  163,  204 
Tetracus,  193 

Thames  Valley,  brick-earths  of,  335 
Theridomyidic,  89,  91,  240 
Theridomys,  91,  192 
TheropithectiSi  231 
Thomas,  Mr  O.,  on  the  fauna  of  S.  E. 

Arabia,  262 ;  on  the  fauna  of  Sipora, 

303 

Thomomys,  366 

ThylacimiS)  39 

Thylacoleo,  37 

Thyropoda,  132 

Tibetan  sub-region,  352 

Tierra  del  Fuego,  140 

Tiger,  272 

Tillodontia,  378 

Timor,  46 

Titanomys,  194 

Titanosaurus,  152 

Titahotheriidce,  170,  377 

Titanotheritim,    in   the   Balkans,  57, 

I7«>»  377 

Tobago,  continental,  137 

Tolypeutes,  94 

Tortoises,  Galapagos,  141 

Tosca,  66 

Toxodon,  82 

Toxodontia,  82 

Toxodontidce,  83 

Toxodontothcrium,  83 

Tragelaphus ,  199,  246 

Tragocerosy  199 

Tragulida,   164,  196,  204,  242;  Ori- 
ental, 281 

TraguluS)  164,  186,  204,  281 

Transition  zone,  361 

Tratratratra,  219 

Tree-porcupines,  136 

Triaulacodus ,  91,  240 

Trichechida,  313 

Triconodon,  51 

Triconodontida:,  51 

Trichosums,  36 

Trichys,  278 

Triclis,  36 

Trigonia,  63 

Trinidad,  continental,  137 

Trilophomys,  238 

Tntylodon,  149 

Trogonidcz,  distribution  of,  129 

Trogontherium ,  333 

Tropical  region,  140 


Trygenycteris,  22  r,  232 

Tucotuco,  90 

Tup aia,  270 

Ttipaiidce,  268,  270 

Tuatera,  8,  63 

Ture,  348 

Tylas,  254 

Ty pother iidce,  84 

Typolherium,  84 

Tyrrhenian  sub-region,  357 

Uintatheriidcz,  174,  378 

Ungulates,  E.  Holarctic,  324;  Neo- 
gaeic,  73;  N.  American,  343  ;  num- 
ber of  African,  241;  Oriental,  278; 
Sonoran,  367 

Uria,  347 

Uromys,  41 

Uronycteris,  42 

Uropsihts,  321 

Urotrichtis,  156,  312,  356 

Ursidce,  their  absence  from  Ethiopian 
Africa,  258;  Holarctic,  321 

Ursus,  202,  274,  318;  absence  of,  in 
Ethiopian  region,  236 ;  ornatus, 


Vampire  bats,  70 

Vandeleuria,  278 

Varanus  priscus,  62 

Vicunas,  74,  136 

Viscachas,  136 

Vishnutherium,  186,  204 

Viverra,    182,    191,    195,    234,    273; 

tangalunga,  47 
Viverricula,  273 
Viverridcz,  African,  234;  an  Old 

World  group,  182;  Oriental,  273 
Voles,  160,  267,  313,  318 

W.  Africa  and  Malaysia,  affinity  of 
faunas,  292 

Wallabies,  36 

Wallace,  Dr,  on  extinction  of  Plistocene 
fauna,  18;  on  zoological  regions,  25; 
on  the  Moluccas,  48 ;  on  Celebes,  48 ; 
on  origin  of  Australian  fauna,  54 ; 
on  isolation  of  Australia,  58 ;  on  S. 
American  fauna,  123 ;  on  connection 
between  S.  America  and  Australia, 
125;  on  W.  Indies,  138;  on  con- 
nection of  S.  and  N.  America,  153  ; 
on  the  past  history  of  Ethiopia, 
255;  on  land  connection  between 
India  and  Africa,  257;  on  Oriental 
sub-regions,  266;  on  relationship 
between  W.  African  and  Malayan 


400 


INDEX. 


faunas,  292  ;  on  Javan  fauna,  302 ; 
on  the  former  union  of  Malta  and 
Sicily  with  Italy,  337 

Wallace's  line,  10 

Walruses,  313 

Wapiti,  315 

Wart-hogs,  256,  268 

Water-buck,  245 

Water-chevrotain,  242 

Water-deer,  356 

Water-shrew,  319 

West  Indian  sub-region,  137 

West  Indies  connecting  N.  and  S. 
America,  139 

Western  Arctogsea,  its  extinct  mam- 
mals, 371 

\Vhite-footed  mice,  88,  342 

Wildebeests,  244 


Wolverene,  312 
Wombats,  38 
Wood-hoopoes,  254 
Wood-rats,  342,  344,  366 

Xenurus,  94 
Xeromys,  40,  277 
Xerus,  195,  197 
Xiphodon,  185,  192,  193 
Xotodon,  83 

Yak,  314 

Zaglossits,  33 

Zapus,  342 

Zebras,  247 

Zoological  realms  and  regions,  25,  27 

Zoster  ops,  254 


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